Morphological and environmental differentiation as prezygotic reproductive barriers between parapatric and allopatric Campanula rotundifolia agg. cytotypes.

BACKGROUND AND AIMS: Reproductive isolation and local establishment are necessary for plant speciation. Polyploidy, the possession of more than two complete chromosome sets, creates a strong postzygotic reproductive barrier between diploid and tetraploid cytotypes. However, this barrier weakens between polyploids (e.g. tetraploids and hexaploids). Reproductive isolation may be enhanced by cytotype morphological and environmental differentiation. Moreover, morphological adaptations to local conditions contribute to plant establishment. However, the relative contributions of ploidy level and the environment to morphology have generally been neglected. Thus, the extent of morphological variation driven by ploidy level and the environment was modelled for diploid, tetraploid and hexaploid cytotypes of Campanula rotundifolia agg. Cytotype distribution was updated, and morphological and environmental differentiation was tested in the presence and absence of natural contact zones. METHODS: Cytotype distribution was assessed from 231 localities in Central Europe, including 48 localities with known chromosome counts, using flow cytometry. Differentiation in environmental niche and morphology was tested for cytotype pairs using discriminant analyses. A structural equation model was used to explore the synergies between cytotype, environment and morphology. KEY RESULTS: Tremendous discrepancies were revealed between the reported and detected cytotype distribution. Neither mixed-ploidy populations nor interploidy hybrids were detected in the contact zones. Diploids had the broadest environmental niche, while hexaploids had the smallest and specialized niche. Hexaploids and spatially isolated cytotype pairs differed morphologically, including allopatric tetraploids. While leaf and shoot morphology were influenced by environmental conditions and polyploidy, flower morphology depended exclusively on the cytotype. CONCLUSIONS: Reproductive isolation mechanisms vary between cytotypes. While diploids and polyploids are isolated postzygotically, the environmental niche shift is essential between higher polyploids. The impact of polyploidy and the environment on plant morphology implies the adaptive potential of polyploids, while the exclusive relationship between flower morphology and cytotype highlights the role of polyploidy in reproductive isolation.

The ecology of palm genomes: repeat-associated genome size expansion is constrained by aridity.

Genome size varies 2400-fold across plants, influencing their evolution through changes in cell size and cell division rates which impact plants' environmental stress tolerance. Repetitive element expansion explains much genome size diversity, and the processes structuring repeat 'communities' are analogous to those structuring ecological communities. However, which environmental stressors influence repeat community dynamics has not yet been examined from an ecological perspective. We measured genome size and leveraged climatic data for 91% of genera within the ecologically diverse palm family (Arecaceae). We then generated genomic repeat profiles for 141 palm species, and analysed repeats using phylogenetically informed linear models to explore relationships between repeat dynamics and environmental factors. We show that palm genome size and repeat 'community' composition are best explained by aridity. Specifically, Ty3-gypsy and TIR elements were more abundant in palm species from wetter environments, which generally had larger genomes, suggesting amplification. By contrast, Ty1-copia and LINE elements were more abundant in drier environments. Our results suggest that water stress inhibits repeat expansion through selection on upper genome size limits. However, elements that may associate with stress-response genes (e.g. Ty1-copia) have amplified in arid-adapted palm species. Overall, we provide novel evidence of climate influencing the assembly of repeat 'communities'.

Nuclear Genome Size in Contrast to Sex Chromosome Number Variability in the Human Bed Bug, Cimex lectularius (Heteroptera: Cimicidae).

The human bed bug Cimex lectularius is one of the most prevalent human ectoparasites in temperate climate zones. The cytogenetic features of this resilient pest include holokinetic chromosomes, special chromosome behavior in meiosis, and numerical variation of chromosomes, where the diploid number ranges from 26 + X1 X2 Y to 26 + X1-20 Y. It is desirable to assess the nuclear DNA content of various cytotypes for a further detailed study of the C. lectularius genome. Detailed knowledge of the DNA content of this parasite could also clarify the origin of additional chromosomes. The average nuclear genome size C. lectularius with 2n = 26 + X1 X2 Y is 2C = 1.94 pg for males and 1.95 pg for females. There is a significant correlation between genome size and the number of chromosomes, but in some specimens with additional chromosomes, nuclear genome size decreases or remains average. Several species used as the internal reference standard were tested for further investigations of genome size in C. lectularius, and the plant Solanum pseudocaspicum turned out to be the most suitable. © 2019 International Society for Advancement of Cytometry.

Ploidy-altered phenotype interacts with local environment and may enhance polyploid establishment in Knautia serpentinicola (Caprifoliaceae).

Whole genome duplication is a key process in plant evolution and has direct phenotypic consequences. However, it remains unclear whether ploidy-related phenotypic changes can significantly alter the fitness of polyploids in nature and thus contribute to establishment of new polyploid mutants in diploid populations. We addressed this question using a unique natural system encompassing a diploid and its sympatric locally established autotetraploid derivative. By setting a common garden experiment with two manipulated environmental factors (presence/absence of serpentine substrate and competition), we tested whether these two locally important factors differently shape the phenotypic response of the two ploidy levels. Tetraploids attained significantly higher values of both above- and below-ground biomass, and root : shoot ratio compared to their diploid progenitors. Tetraploid superiority in vegetative fitness indicators was most prominent when they were cultivated together with a competitor in nutrient-rich nonserpentine substrate. We show that even genetically very closely related diploids and tetraploids can respond differently to key environmental factors. Provided there are sufficient nutrients, tetraploids can be more successful in tolerating interspecific competition than their diploid progenitors. Such superior performance might have provided an adaptive advantage for the newly established tetraploid promoting colonisation of new (micro-)habitats, which was indeed observed at the natural site.

Diversity in genome size and GC content shows adaptive potential in orchids and is closely linked to partial endoreplication, plant life-history traits and climatic conditions.

In angiosperms, genome size and nucleobase composition (GC content) exhibit pronounced variation with possible adaptive consequences. The hyperdiverse orchid family possessing the unique phenomenon of partial endoreplication (PE) provides a great opportunity to search for interactions of both genomic traits with the evolutionary history of the family. Using flow cytometry, we report values of both genomic traits and the type of endoreplication for 149 orchid species and compare these with a suite of life-history traits and climatic niche data using phylogeny-based statistics. The evolution of genomic traits was further studied using the Brownian motion (BM) and Ornstein-Uhlenbeck (OU) models to access their adaptive potential. Pronounced variation in genome size (341-54 878 Mb), and especially in GC content (23.9-50.5%), was detected among orchids. Diversity in both genomic traits was closely related to the type of endoreplication, plant growth form and climatic conditions. GC content was also associated with the type of dormancy. In all tested scenarios, OU models always outperformed BM models. Unparalleled GC content variation was discovered in orchids, setting new limits for plants. Our study indicates that diversity in both genome size and GC content has adaptive consequences and is tightly linked with evolutionary transitions to PE.

Small genome separates native and invasive populations in an ecologically important cosmopolitan grass.

The literature suggests that small genomes promote invasion in plants, but little is known about the interaction of genome size with other traits or about the role of genome size during different phases of the invasion process. By intercontinental comparison of native and invasive populations of the common reed Phragmites australis, we revealed a distinct relationship between genome size and invasiveness at the intraspecific level. Monoploid genome size was the only significant variable that clearly separated the North American native plants from those of European origin. The mean Cx value (the amount of DNA in one chromosome set) for source European native populations was 0.490 ± 0.007 (mean ± SD), for North American invasive 0.506 ± 0.020, and for North American native 0.543 ± 0.021. Relative to native populations, the European populations that successfully invaded North America had a smaller genome that was associated with plant traits favoring invasiveness (long rhizomes, early emerging abundant shoots, resistance to aphid attack, and low C:N ratio). The knowledge that invasive populations within species can be identified based on genome size can be applied to screen potentially invasive populations of Phragmites in other parts of the world where they could grow in mixed stands with native plants, as well as to other plant species with intraspecific variation in invasion potential. Moreover, as small genomes are better equipped to respond to extreme environmental conditions such as drought, the mechanism reported here may represent an emerging driver for future invasions and range expansions.

Sympatric diploid and tetraploid cytotypes of Centaurea stoebe s.l. do not differ in arbuscular mycorrhizal communities and mycorrhizal growth response.

PREMISE OF THE STUDY: Genome duplication is associated with multiple changes at different levels, including interactions with pollinators and herbivores. Yet little is known whether polyploidy may also shape belowground interactions. METHODS: To elucidate potential ploidy-specific interactions with arbuscular mycorrhizal fungi (AMF), we compared mycorrhizal colonization and assembly of AMF communities in roots of diploid and tetraploid Centaurea stoebe s.l. (Asteraceae) co-occurring in a Central European population. In a follow-up greenhouse experiment, we tested inter-cytotype differences in mycorrhizal growth response by combining ploidy, substrate, and inoculation with native AMF in a full-factorial design. KEY RESULTS: All sampled plants were highly colonized by AMF, with the Glomeraceae predominating. AMF-community composition revealed by 454-pyrosequencing reflected the spatial distribution of the hosts, but not their ploidy level or soil characteristics. In the greenhouse experiment, the tetraploids produced more shoot biomass than the diploids did when grown in a more fertile substrate, while no inter-cytotype differences were found in a less fertile substrate. AMF inoculation significantly reduced plant growth and improved P uptake, but its effects did not differ between the cytotypes. CONCLUSIONS: The results do not support our hypotheses that the cytotype structure in a mixed-ploidy population of C. stoebe is mirrored in AMF-community composition and that ploidy-specific fungal communities contribute to cytotype co-existence. Causes and implications of the observed negative growth response to AMF are discussed.

Polyploid species rely on vegetative reproduction more than diploids: a re-examination of the old hypothesis.

Background and Aims: Polyploidy is arguably the single most important genetic mechanism in plant speciation and diversification. It has been repeatedly suggested that polyploids show higher vegetative reproduction than diploids (to by-pass low fertility after the polyploidization), but there are no rigorous tests of it. Methods: Data were analysed by phylogenetic regressions of clonal growth parameters, and vegetative reproduction in culture on the ploidy status of a large set of species (approx. 900) from the Central European Angiosperm flora. Further, correlated evolution of ploidy and clonal traits was examined to determine whether or not polyploidy precedes vegetative reproduction. Key Results: The analyses showed that polyploidy is strongly associated with vegetative reproduction, whereas diploids rely more on seed reproduction. The rate of polyploid speciation is strongly enhanced by the existence of vegetative reproduction (namely extensive lateral spread), whereas the converse is not true. Conclusions: These findings confirm the old hypothesis that polyploids can rely on vegetative reproduction which thus may save many incipient polyploids from extinction. A closer analysis also shows that the sequence of events begins with development of vegetative reproduction, which is then followed by polyploidy. Vegetative reproduction is thus likely to play an important role in polyploid speciation.

Evolutionary dynamics of mixed-ploidy populations in an annual herb: dispersal, local persistence and recurrent origins of polyploids.

Background and Aims: Despite the recent wealth of studies targeted at contact zones of cytotypes in various species, some aspects of polyploid evolution are still poorly understood. This is especially the case for the frequency and success rate of spontaneous neopolyploidization or the temporal dynamics of ploidy coexistence, requiring massive ploidy screening and repeated observations, respectively. To fill this gap, an extensive study of spatio-temporal patterns of ploidy coexistence was initiated in the widespread annual weed Tripleurospermum inodorum (Asteraceae). Methods: DNA flow cytometry along with confirmatory chromosome counts was employed to assess ploidy levels of 11 018 adult individuals and 1263 ex situ germinated seedlings from 1209 Central European populations. The ploidy screening was conducted across three spatial scales and supplemented with observations of temporal development of 37 mixed-ploidy populations. Key Results: The contact zone between the diploid and tetraploid cytotypes has a diffuse, mosaic-like structure enabling common cytotype coexistence from the within-population to the landscape level. A marked difference in monoploid genome size between the two cytotypes enabled the easy distinction of neotetraploid mutants from long-established tetraploids. Neotetraploids were extremely rare (0·03 %) and occurred solitarily. Altogether five ploidy levels (2 x -6 x ) and several aneuploids were discovered; the diversity in nuclear DNA content was highest in early ontogenetic stages (seedlings) and among individuals from mixed-ploidy populations. In spite of profound temporal oscillations in cytotype frequencies in mixed-ploidy populations, both diploids and tetraploids usually persisted up to the last census. Conclusions: Diploids and tetraploids commonly coexist at all spatial scales and exhibit considerable temporal stability in local ploidy mixtures. Mixed-ploidy populations containing fertile triploid hybrids probaby act as effective generators of cytogenetic novelty and may facilitate inter-ploidy gene flow. Neopolyploid mutants were incapable of local establishment.

Ecological differentiation of diploid and polyploid cytotypes of Senecio carniolicus sensu lato (Asteraceae) is stronger in areas of sympatry.

BACKGROUND AND AIMS: Ecological differentiation is recognized as an important factor for polyploid speciation, but little is known regarding whether the ecological niches of cytotypes differ between areas of sympatry and areas where single cytotypes occur (i.e. niche displacement). METHODS: Ecological niches of four groups of Senecio carniolicus sensu lato (s.l.) (western and eastern diploid lineages, tetraploids and hexaploids) were characterized via Landolt indicator values of the accompanying vascular plant species and tested using multivariate and univariate statistics. KEY RESULTS: The four groups of S. carniolicus s.l. were ecologically differentiated mainly with respect to temperature, light and soil (humus content, nutrients, moisture variability). Niche breadths did not differ significantly. In areas of sympatry hexaploids shifted towards sites with higher temperature, less light and higher soil humus content as compared with homoploid sites, whereas diploids and tetraploids shifted in the opposite direction. In heteroploid sites of tetraploids and the western diploid lineage the latter shifted towards sites with lower humus content but higher aeration. CONCLUSIONS: Niche displacement can facilitate the formation of stable contact zones upon secondary contact of polyploids and their lower-ploid ancestors and/or lead to convergence of the cytotypes' niches after they have attained non-overlapping ranges. Niche displacement is essential for understanding ecological consequences of polyploidy.

Species-rich and polyploid-poor: Insights into the evolutionary role of whole-genome duplication from the Cape flora biodiversity hotspot.

PREMISE OF THE STUDY: Whole-genome duplication (WGD) in angiosperms has been hypothesized to be advantageous in unstable environments and/or to increase diversification rates, leading to radiations. Under the first hypothesis, floras in stable environments are predicted to have lower proportions of polyploids than highly, recently disturbed floras, whereas species-rich floras would be expected to have higher than expected proportions of polyploids under the second. The South African Cape flora is used to discriminate between these two hypotheses because it features a hyperdiverse flora predominantly generated by a limited number of radiations (Cape clades), against a backdrop of climatic and geological stability. METHODS: We compiled all known chromosome counts for species in 21 clades present in the Cape (1653 species, including 24 Cape clades), inferred ploidy levels for these species by inspection or derived from the primary literature, and compared Cape to non-Cape ploidy levels in these clades (17,520 species) using G tests. KEY RESULTS: The Cape flora has anomalously low proportions of polyploids compared with global levels. This pattern is consistently observed across nearly half the clades and across global latitudinal gradients, although individual lineages seem to be following different paths to low levels of WGD and to differing degrees. CONCLUSIONS: This pattern shows that the diversity of the Cape flora is the outcome of primarily diploid radiations and supports the hypothesis that WGD may be rare in stable environments.

Challenges of flow-cytometric estimation of nuclear genome size in orchids, a plant group with both whole-genome and progressively partial endoreplication.

Nuclear genome size is an inherited quantitative trait of eukaryotic organisms with both practical and biological consequences. A detailed analysis of major families is a promising approach to fully understand the biological meaning of the extensive variation in genome size in plants. Although Orchidaceae accounts for ∼10% of the angiosperm diversity, the knowledge of patterns and dynamics of their genome size is limited, in part due to difficulties in flow cytometric analyses. Cells in various somatic tissues of orchids undergo extensive endoreplication, either whole-genome or partial, and the G1-phase nuclei with 2C DNA amounts may be lacking, resulting in overestimated genome size values. Interpretation of DNA content histograms is particularly challenging in species with progressively partial endoreplication, in which the ratios between the positions of two neighboring DNA peaks are lower than two. In order to assess distributions of nuclear DNA amounts and identify tissue suitable for reliable estimation of nuclear DNA content, we analyzed six different tissue types in 48 orchid species belonging to all recognized subfamilies. Although traditionally used leaves may provide incorrect C-values, particularly in species with progressively partial endoreplication, young ovaries and pollinaria consistently yield 2C and 1C peaks of their G1-phase nuclei, respectively, and are, therefore, the most suitable parts for genome size studies in orchids. We also provide new DNA C-values for 22 orchid genera and 42 species. Adhering to the proposed methodology would allow for reliable genome size estimates in this largest plant family. Although our research was limited to orchids, the need to find a suitable tissue with dominant 2C peak of G1-phase nuclei applies to all endopolyploid species.

The hidden side of plant invasions: the role of genome size.

The ecological role of genome size in plant biology, biogeography, and morphology has garnered increasing attention as the methods and technology associated with measuring cytological characteristics have become more reliable and accessible. However, how plant genome size influences plant invasions and at what stage in the invasion this influence occurs have been little explored. Several large-scale analyses of published data have yielded valuable interspecific comparisons, but experimental studies that manipulate environmental factors are needed, particularly below the species level, to fully understand the role that genome size plays in plant invasion. In this review, we summarize the available knowledge, discuss the integration of genome size data into invasion research, and suggest how it can be applied to detect and manage invasive species. We also explore how global climate change could exert selective pressures on plant populations with varying genome sizes, thereby increasing the distribution range and invasiveness of some populations while decreasing others. Finally, we outline avenues for future research, including considerations of large-scale studies of intraspecific variation in genome size of invasive populations, testing the interaction of genome size with other factors in macroecological analyses of invasions, as well as the role this trait may play in plant-enemy interactions.

The origin of unique diversity in deglaciated areas: traces of Pleistocene processes in north-European endemics from the Galium pusillum polyploid complex (Rubiaceae).

The role of glacial oscillations in shaping plant diversity has been only rarely addressed in endemics of formerly glaciated areas. The Galium pusillum group represents a rare example of an ecologically diverse and ploidy-variable species complex that exhibits substantial diversity in deglaciated northern Europe. Using AFLP and plastid and nuclear DNA sequences of 67 populations from northern, central, and western Europe with known ecological preferences, we elucidate the evolutionary history of lineages restricted to deglaciated areas and identify the eco-geographic partitioning of their genetic variation. We reveal three distinct endemic northern lineages: (i) diploids from southern Sweden + the British Isles, (ii) tetraploids from southern Scandinavia and the British Isles that show signs of ancient hybridization between the first lineage and populations from unglaciated central Europe, and (iii) tetraploids from Iceland + central Norway. Available evidence supports a stepwise differentiation of these three lineages that started at least before the last glacial maximum by processes of genome duplication, interlineage hybridization and/or allopatric evolution in distinct periglacial refugia. We reject the hypothesis of more recent postglacial speciation. Ecological characteristics of the populations under study only partly reflect genetic variation and suggest broad niches of postglacial colonizers. Despite their largely allopatric modern distributions, the north-European lineages of the G. pusillum group do not show signs of rapid postglacial divergence, in contrast to most other northern endemics. Our study suggests that plants inhabiting deglaciated areas outside the Arctic may exhibit very different evolutionary histories compared with their more thoroughly investigated high-arctic counterparts.

Niche partitioning in arbuscular mycorrhizal communities in temperate grasslands: a lesson from adjacent serpentine and nonserpentine habitats.

Arbuscular mycorrhizal fungi (AMF) represent an important soil microbial group playing a fundamental role in many terrestrial ecosystems. We explored the effects of deterministic (soil characteristics, host plant life stage, neighbouring plant communities) and stochastic processes on AMF colonization, richness and community composition in roots of Knautia arvensis (Dipsacaceae) plants from three serpentine grasslands and adjacent nonserpentine sites. Methodically, the study was based on 454-sequencing of the ITS region of rDNA. In total, we detected 81 molecular taxonomical operational units (MOTUs) belonging to the Glomeromycota. Serpentine character of the site negatively influenced AMF root colonization, similarly as higher Fe concentration. AMF MOTUs richness linearly increased along a pH gradient from 3.5 to 5.8. Contrary, K and Cr soil concentration had a negative influence on AMF MOTUs richness. We also detected a strong relation between neighbouring plant community composition and AMF MOTUs richness. Although spatial distance between the sampled sites (c. 0.3-3 km) contributed to structuring AMF communities in K. arvensis roots, environmental parameters were key factors in this respect. In particular, the composition of AMF communities was shaped by the complex of serpentine conditions, pH and available soil Ni concentration. The composition of AMF communities was also dependent on host plant life stage (vegetative vs. generative). Our study supports the dominance of deterministic factors in structuring AMF communities in heterogeneous environment composed of an edaphic mosaic of serpentine and nonserpentine soils.

Naturalization of central European plants in North America: species traits, habitats, propagule pressure, residence time.

The factors that promote invasive behavior in introduced plant species occur across many scales of biological and ecological organization. Factors that act at relatively small scales, for example, the evolution of biological traits associated with invasiveness, scale up to shape species distributions among different climates and habitats, as well as other characteristics linked to invasion, such as attractiveness for cultivation (and by extension propagule pressure). To identify drivers of invasion it is therefore necessary to disentangle the contribution of multiple factors that are interdependent. To this end, we formulated a conceptual model describing the process of invasion of central European species into North America based on a sequence of "drivers." We then used confirmatory path analysis to test whether the conceptual model is supported by a statistical model inferred from a comprehensive database containing 466 species. The path analysis revealed that naturalization of central European plants in North America, in terms of the number of North American regions invaded, most strongly depends on residence time in the invaded range and the number of habitats occupied by species in their native range. In addition to the confirmatory path analysis, we identified the effects of various biological traits on several important drivers of the conceptualized invasion process. The data supported a model that included indirect effects of biological traits on invasion via their effect on the number of native range habitats occupied and cultivation in the native range. For example, persistent seed banks and longer flowering periods are positively correlated with number of native habitats, while a stress-tolerant life strategy is negatively correlated with native range cultivation. However, the importance of the biological traits is nearly an order of magnitude less than that of the larger scale drivers and highly dependent on the invasion stage (traits were associated only with native range drivers). This suggests that future research should explicitly link biological traits to the different stages of invasion, and that a failure to consider residence time or characteristics of the native range may seriously overestimate the role of biological traits, which, in turn, may result in spurious predictions of plant invasiveness.

Nonadaptive processes governing early stages of polyploid evolution: Insights from a primary contact zone of relict serpentine Knautia arvensis (Caprifoliaceae).

• Premise of the study: Contact zones between polyploids and their diploid progenitors may provide important insights into the mechanisms of sympatric speciation and local adaptation. However, most published studies investigated secondary contact zones where the effects of genome duplication can be confounded by previous independent evolution of currently sympatric cytotypes. We compared genetically close diploid and autotetraploid serpentine cytotypes of Knautia arvensis (Caprifoliaceae) in a primary contact zone and evaluated the role of adaptive and nonadaptive processes for cytotype coexistence.• Methods: DNA flow cytometry was used to determine ploidy distribution at various spatial scales (from across the entire contact zone to microgeographic). Habitat preferences of diploids and polyploids were assessed by comparing vegetation composition of nearby ploidy-uniform sites and by recording plant species immediately surrounding both cytotypes in mixed-ploidy plots.• Key results: Tetraploids considerably outnumbered their diploid progenitors in the contact zone. Both cytotypes were segregated at all investigated spatial scales. This pattern was not driven by ecological shifts, because both diploids and tetraploids inhabited sites with nearly identical vegetation cover. Certain interploidy niche differentiation was indicated only at the smallest spatial scale; ecologically nonadaptive processes were most likely responsible for this difference.• Conclusions: We conclude that a shift in ecological preferences (i.e., the adaptive scenario) is not necessary for the establishment and evolutionary success of autopolyploid derivatives in primary contact zones. Spatial segregation that would support ploidy coexistence can also be achieved by ecologically nonadaptive processes, including the founder effect, limited dispersal ability, intense clonal growth, and triploid block.

Intraspecific ploidy variation: A hidden, minor player in plant-soil-mycorrhizal fungi interactions.

PREMISE OF THE STUDY: Genome duplication and arbuscular mycorrhizal (AM) symbiosis are ubiquitous in angiosperms. While the significance of each of these phenomena separately has been intensively studied, their interaction remains to be understood. METHODS: Three diploid and three hexaploid populations of Aster amellus (Asteraceae) were characterized in terms of the soil conditions in situ and mycorrhizal root colonization. In a greenhouse experiment, the effects of ploidy level, substrate conditions, and AM fungi on plant performance were then separated by growing noninoculated plants or plants inoculated with AM fungi in substrates native to either the diploids or hexaploids. KEY RESULTS: The diploids inhabited nutritionally richer sites but did not differ from hexaploid plants in the level of mycorrhizal root colonization in situ. In the experiment, hexaploids generally performed better than the diploids. This intercytotype growth difference was enhanced by soil fertility, with hexaploids benefiting more from nutritionally richer substrate than the diploids. AM inoculation was crucial for plant growth and phosphorus uptake. The interaction between ploidy level and AM inoculation significantly influenced only dry mass of roots, phosphorus concentrations in shoot biomass, and the length of the extraradical mycelium in the nonsterile substrates. CONCLUSIONS: Our results support the idea that polyploidy can affect the mycorrhizal growth response of host plants. Nevertheless, the effects of the interaction between ploidy and inoculation were weaker than the main effects of these factors.

Ploidy frequencies in plants with ploidy heterogeneity: fitting a general gametic model to empirical population data.

Genome duplication (polyploidy) is a recurrent evolutionary process in plants, often conferring instant reproductive isolation and thus potentially leading to speciation. Outcome of the process is often seen in the field as different cytotypes co-occur in many plant populations. Failure of meiotic reduction during gametogenesis is widely acknowledged to be the main mode of polyploid formation. To get insight into its role in the dynamics of polyploidy generation under natural conditions, and coexistence of several ploidy levels, we developed a general gametic model for diploid-polyploid systems. This model predicts equilibrium ploidy frequencies as functions of several parameters, namely the unreduced gamete proportions and fertilities of higher ploidy plants. We used data on field ploidy frequencies for 39 presumably autopolyploid plant species/populations to infer numerical values of the model parameters (either analytically or using an optimization procedure). With the exception of a few species, the model fit was very high. The estimated proportions of unreduced gametes (median of 0.0089) matched published estimates well. Our results imply that conditions for cytotype coexistence in natural populations are likely to be less restrictive than previously assumed. In addition, rather simple models show sufficiently rich behaviour to explain the prevalence of polyploids among flowering plants.

Ploidy-specific symbiotic interactions: divergence of mycorrhizal fungi between cytotypes of the Gymnadenia conopsea group (Orchidaceae).

Polyploidy is widely recognized as a major mechanism of sympatric speciation in plants, yet little is known about its effects on interactions with other organisms. Mycorrhizal fungi are among the most common plant symbionts and play an important role in plant nutrient supply. It remains to be understood whether mycorrhizal associations of ploidy-variable plants can be ploidy-specific. We examined mycorrhizal associations in three cytotypes (2x, 3x, 4x) of the Gymnadenia conopsea group (Orchidaceae), involving G. conopsea s.s. and G. densiflora, at different spatial scales and during different ontogenetic stages. We analysed: adults from mixed- and single-ploidy populations at a regional scale; closely spaced adults within a mixed-ploidy site; and mycorrhizal seedlings. All Gymnadenia cytotypes associated mainly with saprotrophic Tulasnellaceae (Basidiomycota). Nonetheless, both adults and seedlings of diploids and their autotetraploid derivatives significantly differed in the identity of their mycorrhizal symbionts. Interploidy segregation of mycorrhizal symbionts was most pronounced within a site with closely spaced adults. This study provides the first evidence that polyploidization of a plant species can be associated with a shift in mycorrhizal symbionts. This divergence may contribute to niche partitioning and facilitate establishment and co-existence of different cytotypes.