Ulla Carlsson-Granér

ANTHER-SMUT DISEASE IN SILENE DIOICA: VARIATION IN SUSCEPTIBILITY AMONG GENOTYPES AND POPULATIONS, AND PATTERNS OF DISEASE WITHIN POPULATIONS

In an archipelago in northern Sweden, populations of the perennial, dioecious, and insect‐pollinated herb Silene dioica are commonly infected by the sterilising anther‐smut fungus Microbotryum violaceum. The results from transplantation and inoculation experiments in this study show that variation between populations in the incidence of disease may partly be due to variation in resistance among populations. In the transplantation experiment in which plants were naturally exposed to the fungus, disease levels varied greatly among transplants from three healthy populations when transplanted to three diseased populations in the archipelago. Clear genotypic differences in susceptibility among 25 replicated genotypes of the host plants were found when inoculated manually with two different isolates. Susceptibility varied between 0‐90%, but the two isolates used did not differ in inoculation success. The results also suggest a geographical structuring in resistance of the host and virulence of the fungus. First, disease levels among experimental plants from two of the disease‐free populations of S. dioica (originating from inner and outer archipelago, respectively), were high when transplanted to a diseased population nearby, but low when transplanted far away. Second, regardless of origin, plants from all healthy populations became diseased in the diseased experimental populations located in the middle part of the archipelago. Due to isostatic land upheaval in the studied archipelago, there is a vertical age‐axis within islands such that the highest point on an island is the oldest. Since this may affect the demography of the host, disease spread, and the dynamics of disease, spatial patterns in adult and seedling densities, and disease and spore deposition along the age‐axis were studied within three diseased populations. A low incidence of disease was found in the young, low and old, high parts of the populations and a high incidence in the vertical, middle parts of the populations. The higher disease incidence in the middle part compared with the lower part of the population may reflect less disturbance and an increased probability of disease with age.

Variation in Breeding System Traits within and among Populations of Microbotryum violaceum on Silene dioica

Breeding systems exert profound effects on the amount and distribution of genetic diversity within and among populations. Knowledge of breeding systems is also important for understanding dynamics between coevolving organisms, e.g., pathogen‐host interactions. Here we study the breeding system of the obligate anther smut Microbotryum violaceum on Silene dioica. Microbotryum violaceum is capable of both inbreeding and outcrossing, but several recent studies on other host races have indicated that automixis via intrapromycelial mating is the predominant breeding system. Compared with conjugations between cells from different meioses, automixis results in slower loss of heterozygosity and faster production of infectious hypha. However, high rates of intrapromycelial matings have been suggested to invoke a fitness cost due to production of fewer infectious dikaryons. Working with single strains under standardized laboratory conditions, we studied traits that could influence the distribution of genetic variability and pathogen fitness. We found that intrapromycelial mating is the dominant conjugation form for M. violaceum var. dioica but that the breeding system varies, partly because of genetic differences, both within and among populations. Further, we did not find the predicted fitness reduction for intrapromycelial matings, suggesting that intrapromycelial mating is a highly favorable breeding system for M. violaceum.

Host resistance and pathogen infectivity in host populations with varying connectivity

Theory predicts that hosts and pathogens will evolve higher resistance and aggressiveness in systems where populations are spatially connected than in situations in which populations are isolated and dispersal is more local. In a large cross‐inoculation experiment we surveyed patterns of host resistance and pathogen infectivity in anther‐smut diseased Viscaria alpina populations from three contrasting areas where populations range from continuous, through patchy but spatially connected to highly isolated demes. In agreement with theory, isolated populations of V. alpina were more susceptible on average than either patchily distributed or continuous populations. While increased dispersal in connected systems increases disease spread, it may also increase host gene flow and the potential for greater host resistance to evolve. In the Viscaria–Microbotryum system, pathogen infectivity mirrored patterns of host resistance with strains from the isolated populations being the least infective and strains from the more resistant continuous populations being the most infective on average, suggesting that high resistance selects for high infectivity. To our knowledge this study is the first to characterize the impacts of varying spatial connectivity on patterns of host resistance and pathogen infectivity in a natural system.

Patterns of disease and host resistance in spatially structured systems

We use data from species of the anther-smut fungi and the host plants Lychnis alpina and Silene dioica to show that spatial structuring at different scales can influence patterns of disease and host resistance. Patterns of disease and host resistance were surveyed in an archipelago subject to land-uplift where populations of S. dioica constitute an age-structured metapopulation, and in three contrasting areas within the mainland range of L. alpina, where population distributions range from continuous, through patchy but spatially connected to highly isolated demes. In S. dioica, disease levels depend on the age, size and density of local patches and populations. Disease is most predictably found in larger dense host patches and populations of intermediate age, and more frequently goes extinct in small old populations. The rate of local disease spread is affected by the level of host resistance; S. dioica populations showing an increase in disease over time are more susceptible than populations where the disease has remained at low levels. Among-population variation in resistance is driven by founding events and populations remain differentiated due to limited gene flow between islands. As observed in the L. alpina system, when populations are more connected, a greater fraction of populations have disease present. Results from a simulation model argue that, while increased dispersal in connected systems can increase disease spread, it may also favour selection of host resistance which ultimately reduces disease levels within populations. This could explain the observed lower disease prevalence in L. alpina in regions where populations are more continuous.

Geographical pattern in the response of the arctic-alpine Silene suecica (Cariophyllaceae) to the interaction between water availability and photoperiod

We hypothesized a geographical pattern of the plant performance (seedling development, biomass production, relative water content and chlorophyll content) as a result of response to the interaction between photoperiod and water availability in populations of the arctic-alpine Silene suecica from different latitudes, thus experiencing different photoperiods during the growing season. Particularly, we expected a lower drought sensitivity in northern compared to southern populations as a consequence of harsher conditions experienced by the northern populations in terms of water availability. The experiment was carried out under common garden conditions, manipulating the water availability (wet and dry) and the photoperiod (21 and 16xa0h). We found an interaction between photoperiod and water availability on plant height, leaves, growth, biomass and total chlorophyll. However, the photoperiod neither counteracted nor intensified the effect of drought. Plants exposed to drought compensated for decreasing water availability by reducing their shoot growth. Changes in the chlorophyll content and chlorophyll a/b ratio were observed. Northern populations showed a higher basal growth performance and a greater response to the changed water regime (from wet to dry) than the southern populations. Southern populations showed a reduced ability to respond to drought, but their low basal performance may be advantageous under low water availability, avoiding water loss. In contrast, northern populations showed a stronger plastic response that limited the negative effects of reduced water availability. This study highlights the possibility that the plant response to environmental constraints (specifically water availability) may follow a geographical pattern.

Seed dormancy and longevity in subarctic and alpine populations of Silene suecica

Despite the strong environmental control of seed dormancy and longevity, their changes along latitudes are poorly understood. The aim of this study was to assess seed dormancy and longevity in different populations across the distribution of the arctic–alpine plant Silene suecica. Seeds of seven populations collected from alpine (Spain, Italy, Scotland) and subarctic (Sweden, Norway) populations were incubated at four temperature regimes and five cold stratification intervals for germination and dormancy testing. Seed longevity was studied by exposing seeds to controlled ageing (45xa0°C, 60% RH) and regularly sampled for germination. Fresh seeds of S. suecica germinated at warm temperature (20/15xa0°C) and more in subarctic (80–100%) compared to alpine (20–50%) populations showed a negative correlation with autumn temperature (i.e., post-dispersal period). Seed germination increased after cold stratification in all populations, with different percentages (30–100%). Similarly, there was a large variation of seed longevity (p50xa0=xa012–32xa0days), with seeds from the wettest locations showing faster deterioration rate. Subarctic populations of S. suecica were less dormant, showing a warmer suitable temperature range for germination, and a higher germinability than alpine populations. Germination and dormancy were driven by an interplay of geographical and climatic factors, with alpine and warm versus subarctic and cool autumn conditions, eliciting a decrease and an increase of emergence, respectively. Germination and dormancy patterns typically found in alpine habitats may not be found in the arctic.