Comparative seed germination traits in alpine and subalpine grasslands: higher elevations are associated with warmer germination temperatures.

Seed germination traits in alpine grasslands are poorly understood, despite the sensitivity of these communities to climate change. We hypothesise that germination traits predict species occurrence along the alpine-subalpine elevation gradient. Phylogenetic comparative analyses were performed using fresh seeds of 22 species from alpine and subalpine grasslands (1600-2400 m) of the Cantabrian Mountains, Spain (43° N, 5° W). Laboratory experiments were conducted to characterise germinability, optimum germination temperature and effect of cold and warm stratification on dormancy breaking. Variability in these traits was reduced by phylogenetic principal component analysis (phyl.PCA). Phylogenetic generalised least squares regression (PGLS) was used to fit a model in which species average elevation was predicted from their position on the PCA axes. Most subalpine species germinated in snow-like conditions, whereas most alpine species needed accumulation of warm temperatures. Phylogenetic signal was low. PCA1 ordered species according to overall germinability, whilst PCA2 ordered them according to preference for warm or cold germination. PCA2 significantly predicted species occurrence in the alpine-subalpine gradient, as higher elevation species tended to have warmer germination preferences. Our results show that germination traits in high-mountain grasslands are closely linked to the alpine-subalpine gradient. Alpine species, especially those from stripped and wind-edge communities, prefer warmer germination niches, suggesting that summer emergence prevents frost damage during seedling establishment. In contrast, alpine snowfield and subalpine grassland plants have cold germination niches, indicating that winter emergence may occur under snow to avoid drought stress.

Fernández-Pascual E ，Jiménez-Alfaro B ，Bueno Á 《-》

Seed dormancy and germination changes of snowbed species under climate warming: the role of pre- and post-dispersal temperatures.

Climate warming has major impacts on seed germination of several alpine species, hence on their regeneration capacity. Most studies have investigated the effects of warming after seed dispersal, and little is known about the effects a warmer parental environment may have on germination and dormancy of the seed progeny. Nevertheless, temperatures during seed development and maturation could alter the state of dormancy, affecting the timing of emergence and seedling survival. Here, the interplay between pre- and post-dispersal temperatures driving seed dormancy release and germination requirements of alpine plants were investigated. Three plant species inhabiting alpine snowbeds were exposed to an artificial warming treatment (i.e. +1·5 K) and to natural conditions in the field. Seeds produced were exposed to six different periods of cold stratification (0, 2, 4, 8, 12 and 20 weeks at 0 °C), followed by four incubation temperatures (5, 10, 15 and 20 °C) for germination testing. A warmer parental environment produced either no or a significant increase in germination, depending on the duration of cold stratification, incubation temperatures and their interaction. In contrast, the speed of germination was less sensitive to changes in the parental environment. Moreover, the effects of warming appeared to be linked to the level of (physiological) seed dormancy, with deeper dormant species showing major changes in response to incubation temperatures and less dormant species in response to cold stratification periods. Plants developed under warmer climates will produce seeds with changed germination responses to temperature and/or cold stratification, but the extent of these changes across species could be driven by seed dormancy traits. Transgenerational plastic adjustments of seed germination and dormancy shown here may result from increased seed viability, reduced primary and secondary dormancy state, or both, and may play a crucial role in future plant adaptation to climate change.

Bernareggi G ，Carbognani M ，Mondoni A ，Petraglia A ... -  《-》

Local climate explains degree of seed dormancy in Hypericum elodes L. (Hypericaceae).

Seed dormancy and germination characteristics may vary within species in response to several factors. Knowledge of such variation is crucial to understand plant evolution and adaptation to environmental changes. We examined the correlation of climate and population genetic differentiation (ISSR) with primary seed dormancy and germination behaviour in populations of the Atlantic-European soft-water pool specialist Hypericum elodes. Primary dormancy was measured by analysing seed germination response of fresh seeds and after various periods of cold stratification. Laboratory germination experiments revealed that the single most important factor for promoting germination was cold stratification prior to placing at the germination temperature. However, in agreement with their weaker primary dormancy, the seeds germinated well when fresh, and the benefit of cold stratification was more relaxed for the southern populations. Seeds of all populations demonstrated a near absolute requirement for a light and alternating temperature regime in order to germinate. The promoting effect of alternating temperatures was particularly effective at warm temperatures (mean 20 °C) but not at cool temperatures. Whilst seed germination requirements were similar among populations, the degree of primary dormancy varied considerably and was not associated with population genetic differentiation. Primary dormancy degree was instead associated with local climate: higher temperature in summer and rainfall in winter predicted weak and rapid loss of dormancy. These results suggest that seed maturation environment may play a substantial role in explaining the degree of dormancy in H. elodes, highlighting that physiological dormancy can be modulated by local climate.

Carta A ，Probert R ，Puglia G ，Peruzzi L ，Bedini G ... -  《-》

Effects of Autumn and Spring Heat Waves on Seed Germination of High Mountain Plants.

Alpine plants are considered to be particularly vulnerable to climate change and related extreme episodes, such as heat waves. Despite growing interest in the impact of heat waves on alpine plants, knowledge about their effects on regeneration is still fragmentary. Recruitment from seeds will be crucial for the successful migration and survival of these species and will play a key role in their future adaptation to climate change. In this study, we assessed the impacts of heat waves on the seed germination of 53 high mountain plants from the Northern Apennines (Italy). The seeds were exposed to laboratory simulations of three seasonal temperature treatments, derived from real data recorded at a meteorological station near the species growing site, which included two heat wave episodes that occurred both in spring 2003 and in autumn 2011. Moreover, to consider the effect of increasing drought conditions related to heat waves, seed germination was also investigated under four different water potentials. In the absence of heat waves, seed germination mainly occurred in spring, after seeds had experienced autumn and winter seasons. However, heat waves resulted in a significant increase of spring germination in c. 30% of the species and elicited autumn germination in 50%. When heat waves were coupled with drought, seed germination decreased in all species, but did not stop completely. Our results suggest that in the future, heat waves will affect the germination phenology of alpine plants, especially conditionally dormant and strictly cold-adapted chorotypes, by shifting the emergence time from spring to autumn and by increasing the proportion of emerged seedlings. The detrimental effects of heat waves on recruitment success is less likely to be due to the inhibition of seed germination per se, but rather due to seedling survival in seasons, and temperature and water conditions that they are not used to experiencing. Changes in the proportion and timing of emergence suggest that there may be major implications for future plant population size and structure.

Orsenigo S ，Abeli T ，Rossi G ，Bonasoni P ，Pasquaretta C ，Gandini M ，Mondoni A ... -  《PLoS One》

Effects of the duration of cold stratification on early life stages of the Mediterranean alpine plant Silene ciliata.

Cold stratification provided by snow cover is essential to break seed dormancy in many alpine plant species. The forecast reduction in snow precipitation and snow cover duration in most temperate mountains as a result of global warming could threaten alpine plant populations, especially those at the edge of their species distribution, by altering the dynamics of early life stages. We simulated some effects of a reduction in the snow cover period by manipulating the duration of cold stratification in seeds of Silene ciliata, a Mediterranean alpine specialist. Seeds from three populations distributed along an altitudinal gradient were exposed to different periods of cold stratification (2, 4 and 6 months) in the laboratory and then moved to common garden conditions in a greenhouse. The duration of the cold stratification treatment and population origin significantly affected seed emergence percentage, emergence rate and seedling size, but not the number of seedling leaves. The 6-month and 4-month cold stratification treatments produced higher emergence percentages and faster emergence rates than seeds without cold stratification treatment. No significant cold stratification duration x seed population origin interactions were found, thus differential sensitivity to cold stratification along elevation is not supported.

García-Fernández A ，Escudero A ，Lara-Romero C ，Iriondo JM ... -  《-》