Season-dependent effect of cleistogamy in Lamium amplexicaule: flower type origin versus inbreeding status.

PREMISE: Inbreeding depression is traditionally considered a major factor favoring outcrossing in hermaphrodites. Recent experiments have shown that environmental conditions can influence the magnitude of inbreeding depression, but their relevance in natural populations is unclear. To investigate this, we studied the cleistogamous species Lamium amplexicaule, an annual species with individuals experiencing either spring or autumn environmental conditions. In this species, the proportion of cleistogamous/chasmogamous flowers changes according to seasonal cues (e.g., temperature, photoperiod). Our hypothesis was that the plasticity of cleistogamy is an adaptation to seasonal fitness variation in different flower progenies. METHODS: To test this hypothesis, we produced the three possible types of progenies through controlled crosses: (1) selfed progeny from cleistogamous flowers; (2) chasmogamous selfed progeny; and (3) chasmogamous outcrossed progeny. Progeny fitness was then measured in a common garden in the two reproduction seasons (autumn and spring). RESULTS: The results showed that season had a major impact on fitness. The fitness of the different progeny types changed across seasons, indicating that the effect of cleistogamy on progeny performance is season-dependent, consistent with a previous study in a similar environment. Surprisingly, the flower from which the progeny issued (cleistogamous or chasmogamous) had more impact on fitness than the inbred status of the progeny (selfed versus outcrossed), suggesting a potential role of epigenetic processes. CONCLUSIONS: The observed patterns of variation were not consistent either with adaptation to environment-dependent inbreeding depression or to variation in resource availability, but were possibly consistent with adaptation to seasonal pollinator activity.

Evolutionary potential of a widespread clonal grass under changing climate.

Adaptive responses are probably the most effective long-term responses of populations to climate change, but they require sufficient evolutionary potential upon which selection can act. This requires high genetic variance for the traits under selection and low antagonizing genetic covariances between the different traits. Evolutionary potential estimates are still scarce for long-lived, clonal plants, although these species are predicted to dominate the landscape with climate change. We studied the evolutionary potential of a perennial grass, Festuca rubra, in western Norway, in two controlled environments corresponding to extreme environments in natural populations: cold-dry and warm-wet, the latter being consistent with the climatic predictions for the country. We estimated genetic variances, covariances, selection gradients and response to selection for a wide range of growth, resource acquisition and physiological traits, and compared their estimates between the environments. We showed that the evolutionary potential of F. rubra is high in both environments, and genetic covariances define one main direction along which selection can act with relatively few constraints to selection. The observed response to selection at present is not sufficient to produce genotypes adapted to the predicted climate change under a simple, space for time substitution model. However, the current populations contain genotypes which are pre-adapted to the new climate, especially for growth and resource acquisition traits. Overall, these results suggest that the present populations of the long-lived clonal plant may have sufficient evolutionary potential to withstand long-term climate changes through adaptive responses.

Adaptive differentiation of Festuca rubra along a climate gradient revealed by molecular markers and quantitative traits.

Species response to climate change is influenced by predictable (selective) and unpredictable (random) evolutionary processes. To understand how climate change will affect present-day species, it is necessary to assess their adaptive potential and distinguish it from the effects of random processes. This will allow predicting how different genotypes will respond to forecasted environmental change. Space for time substitution experiments are an elegant way to test the response of present day populations to climate variation in real time. Here we assess neutral and putatively adaptive variation in 11 populations of Festuca rubra situated along crossed gradients of temperature and moisture using molecular markers and phenotypic measurements, respectively. By comparing population differentiation in putatively neutral molecular markers and phenotypic traits (QST-FST comparisons), we show the existence of adaptive differentiation in phenotypic traits and their plasticity across the climatic gradient. The observed patterns of differentiation are due to the high genotypic and phenotypic differentiation of the populations from the coldest (and wettest) environment. Finally, we observe statistically significant covariation between markers and phenotypic traits, which is likely caused by isolation by adaptation. These results contribute to a better understanding of the current adaptation and evolutionary potential to face climate change of a widespread species. They can also be extrapolated to understand how the studied populations will adjust to upcoming climate change without going through the lengthy process of phenotyping.

Isolation and characterization of microsatellite markers for the cleistogamous species Lamium amplexicaule (Lamiaceae).

PREMISE OF THE STUDY: Lamium amplexicaule is a cleistogamous plant that produces both closed flowers (obligately self-pollinated) and open flowers (potentially outcrossed). The conditions for the maintenance of such a mating system depend on the outcrossing rate of the open flowers, which can be estimated using neutral microsatellite markers. • METHODS AND RESULTS: Forty primer pairs corresponding to microsatellite motifs obtained by coupling multiplex microsatellite enrichment and next-generation sequencing were tested. Thirteen primers amplified with satisfying results. The polymorphism of these markers was studied in four French populations. Allele number varied from one to eight per locus and per population. Heterozygosity levels were significantly lower than those expected under Hardy-Weinberg equilibrium. • CONCLUSIONS: Our results are consistent with a partial self-fertilization pattern. These markers will be used to estimate the outcrossing rate as well as population differentiation in L. amplexicaule.