Effects of experimentally imposed climate scenarios on flowering phenology and flower production of subarctic bog species

Climate scenarios for high‐latitude areas predict not only increased summer temperatures, but also larger variation in snowfall and winter temperatures. By using open‐top chambers, we experimentally manipulated both summer temperatures and winter and spring snow accumulations and temperatures independently in a blanket bog in subarctic Sweden, yielding six climate scenarios. We studied the effects of these scenarios on flowering phenology and flower production of Andromeda polifolia (woody evergreen) and Rubus chamaemorus (perennial herb) during 2 years. The second year of our study (2002) was characterized by unusually high spring and early summer temperatures. Our winter manipulations led to consistent increases in winter snow cover. As a result, average and minimum air and soil temperatures in the high snow cover treatments were higher than in the winter ambient treatments, whereas temperature fluctuations were smaller. Spring warming resulted in higher average, minimum, and maximum soil temperatures. Summer warming led to higher air and soil temperatures in mid‐summer (June–July), but not in late summer (August–September). The unusually high temperatures in 2002 advanced the median flowering date by 2 weeks for both species in all treatments. Superimposed on this effect, we found that for both Andromeda and Rubus, all our climate treatments (except summer warming for Rubus) advanced flowering by 1–4 days. The total flower production of both species showed a more or less similar response: flower production in the warm year 2002 exceeded that in 2001 by far. However, in both species flower production was only stimulated by the spring‐warming treatments. Our results show that the reproductive ecology of both species is very responsive to climate change but this response is very dependent on specific climate events, especially those that occur in winter and spring. This suggests that high‐latitude climate change experiments should focus more on winter and spring events than has been the case so far.     

Three-dimensional soil heterogeneity modulates responses of grassland mesocosms to an experimentally imposed drought extreme

Heterogeneity is an intrinsic characteristic of soils, which regulates plant diversity and ecosystem functioning. However, whether soil heterogeneity also modulates responses of plant communities to climate change, including climate extremes, remains largely an open question. Here, we explore responses of plant communities to drought extremes across four levels of spatial soil heterogeneity, with cell sizes varying from very small to very large, i.e. 0, 12, 24 and 48 cm. These were created in mesocosms by alternating nutrient-rich and nutrient-poor substrate in three dimensions. A seed mixture of 24 grassland species was evenly sown on each mesocosm in spring. In late summer, a three-week drought was imposed with a rainout shelter. During the drought, soil water content at the mesocosm scale decreased more at intermediate (12 and 24 cm) than at small or large (0 and 48 cm) cell sizes, which was reflected in increased senescence and drought-induced heat stress. These responses could be traced to greater plant biomass coupled with higher water demand at those intermediate cell sizes, likely related to between-cell access to nutrients and effects of diversity and community composition. Our results indicate that soil heterogeneity can modulate the impact of drought extremes on plant communities, though more research is needed on the transition between intermediate and extreme cell sizes, where heterogeneity effects seem to change most. We propose that soil heterogeneity be considered more explicitly in studies of changing precipitation regimes.     

Site-specific factors influence the richness and phenology of snowbed plants in the Pyrenees

Although the timing of snowmelt and growth temperatures appear to be the main factors that influence the species richness and phenology of snowbed plants, site-specific characteristics may also play a role in modifying the effects of the timing of snowmelt and temperature. In this study, the effects of site-specific factors (microtopography and snow origin) on species richness and plant phenology were evaluated in 72 plots in two snowbeds in the Andorran Pyrenees. Snowmelt patterns influenced the spatial distribution of species richness and abundance. Site-specific factors had significant effects on the responses of species (shortening or lengthening the duration of the phenophase) and on the extent to which the timing of snowmelt influenced leaf expansion and flowering. Notably, the highest rates of leaf expansion occurred on late snowmelt isoclines, where, nevertheless, the time taken to reach peak flowering was significantly longer than on the early snowmelt isoclines. The results of this study highlight the fact that, in addition to the effects of interannual variability in climate, site-specific factors have a significant effect on the phenology and reproductive success of the commonest plants in the snowbed communities of the Pyrenees.     

Effects of nutrient amendments on modular growth,flowering effort and reproduction of snowbed plants

Background: There have been few studies on the effects on alpine flora and vegetation of an increase in nutrient availability. Two main potential sources of increased nutrient availability in alpine ecosystems are enhanced mineralisation caused by climate warming and atmospheric nitrogen deposition. Little is known on how life history traits of different species are related at individual and community levels to enhanced nutrient availability.

Aims: We investigated the effects of nitrogen and phosphorus addition on the modular growth, flowering and germination of four species: two snowbed specialists and two alpine generalists.

Methods: We established in a late snowbed at the Gavia Pass, Italian Alps, a factorial experiment with four fertiliser treatments (N, P, Low N+P, and High N+P) and an unfertilised control. Nutrients were applied in 2003–2006 and we recorded the number of modules per individuals, total number of flowers, number of flowers per flowering module, and calculated the percentage of flowering modules in 2005 and 2006, and counted the number of seedlings in 2006.

Results: The modular growth and flowering effort of the species appeared to be co-limited by N and P. The alpine generalists showed greater responses to fertilization in vegetative growth and flowering compared with snowbed specialists. The number of seedlings recorded indicated species-specific responses to nutrient addition.

Conclusions: Our results suggest that enhanced nutrient availability stimulates modular growth and flowering effort in responsive alpine generalist species. This, in turn, could lead to an increase in their abundance, leading to changes in community structure in snowbeds with enhanced nutrient availability.     

Meta-analysis of phenotypic selection on flowering phenology suggests that early flowering plants are favoured

Flowering times of plants are important life-history components and it has previously been hypothesized that flowering phenologies may be currently subject to natural selection or be selectively neutral. In this study we reviewed the evidence for phenotypic selection acting on flowering phenology using ordinary and phylogenetic meta-analysis. Phenotypic selection exists when a phenotypic trait co-varies with fitness; therefore, we looked for studies reporting an association between two components of flowering phenology (flowering time or flowering synchrony) with fitness. Data sets comprising 87 and 18 plant species were then used to assess the incidence and strength of phenotypic selection on flowering time and flowering synchrony, respectively. The influence of dependence on pollinators, the duration of the reproductive event, latitude and plant longevity as moderators of selection were also explored. Our results suggest that selection favours early flowering plants, but the strength of selection is influenced by latitude, with selection being stronger in temperate environments. However, there is no consistent pattern of selection on flowering synchrony. Our study demonstrates that phenotypic selection on flowering time is consistent and relatively strong, in contrast to previous hypotheses of selective neutrality, and has implications for the evolution of temperate floras under global climate change.     

Effects of nitrogen and phosphorus supply on growth and flowering phenology of the snowbed forb Gnaphalium supinum L.

The warming-induced increase in nutrient mineralization and the further increase in atmospheric nitrogen depositions raise the topic of whether and how alpine plants will react to enhanced nutrient availability. Despite several studies have shown the effects of fertilization on primary production of alpine plants, only few studies have considered the influences of nutrients on reproduction. Here, we investigated the effects of nitrogen (N) and phosphorus (P) amendments on cover, number of ramets, flowering effort and phenological timing of Gnaphalium supinum, an arctic-alpine widespread snowbed species. We set up an experimental design with four fertilization treatments (low N, P without additional N, low N + P, and high N + P) and an unfertilized control for three years (2003–2005), within a late snowbed located in the Italian Alps (Gavia Pass, 2700 m a.s.l.). The cover of Gnaphalium supinum was recorded at the peak of the aboveground biomass development in the three years, while the temporal dynamic of ramet density and reproductive phenophases were monitored during the 2005 growing season. The clonal growth of G. supinum resulted to be co-limited by N and P, while the flowering effort was stimulated by P. Flowering date was advanced by P supply, while N alone did not show any significant effect on phenology. In a warming scenario, with a predicted increase in N and P availability by nutrient mineralization and atmospheric deposition, this species should probably experience some benefits for its growth and reproduction if not limited by other factors such as the length of the growing season or interspecific competition.     

Short-term effects of burn season on flowering phenology of savanna plants

We examined the effect of season of burn on flowering phenology of groundlayer species, in the year following burns, in a mesic-sand Midwestern oak savanna. Burn treatments were fall, early-season, growing-season, late-season, and 1 or 5 years after a prior early-season wildfire. For these treatments, we compared the number of flowering stems and of flowers for species overall, for the 20 most prolifically flowering species, as well as for species grouped by flowering phenoperiods, and by growth form. Growing-season burn had a significant negative effect on number of flowering stems and total number of flowers. This effect occurred when either the burn occurred during the flowering season or during the season prior to the flowering phenoperiod. Tradescantia ohiensis showed expedited flowering and Phlox pilosa showed delayed flowering in response to early-season burning. Flowering of early shrubs was reduced by the previous fall and early-spring fires, while flowering of mid-season blooming shrubs was reduced by the early- and growing-season burns. Vaccinium and Gaylussacia, early-flowering shrubs, produced fewer flowers 1 year after than 5 years after an early-season burn. Arabis lyrata showed reduced flowering from the early-season burn. We also found four instances where the early-spring burn effect on flowering was more severe than the fall burn effect, suggesting that many frequent early-season burns may be deleterious to flowering and reproduction of some species. Burns occurring too frequently in the same season could negatively affect future flowering and reproduction of these plant species.     

Responses of leafing phenology and photosynthesis to soil warming in forest-floor plants

Phenological and physiological responses of plants to climate change are key issues to understand the global change impact on ecosystems. To evaluate the species-specific responses, a soil-warming experiment was conducted for seven understory species having various leaf habits in a deciduous forest, northern Japan; one evergreen shrub, one semi-evergreen fern, one summer-deciduous shrub, and four summer-green herbs. Soil temperature in the warming plots was electrically maintained 5 °C higher than control plots. Responses of leafing phenology highly varied among species: new leaf emergence of the evergreen shrub was delayed; senescence of overwintering leaves of the semi-evergreen fern was accelerated resulting in the shift to deciduousness; leaf shedding of the summer-deciduous shrub was accelerated. Among four summer-green species, only an earliest leaf-out species advanced growth initiation, but the period of growth season was not changed. Physiological responses to soil warming were also highly species-specific: the warming treatment increased the photosynthetic activity of the summer-deciduous shrub and one summer-green species, decreased that of the semi-evergreen fern, while other species did not show any changes in photosynthetic traits. Totally, the soil warming impacts on understory plants was apparent in spring. It was suggested that modification of snow conditions is important issue especially for plants with overwintering leaves. Responses of understory vegetation to climate change may highly vary depending on the composition of leaf habits in the cool-temperate forests.     

荒漠短命植物地上和地下生产力对极端干旱和降水的响应

在古尔班通古特沙漠南缘沙垄4个坡位和坡向,设置减少65%和增加65%生长季降水量以模拟极端干旱和极端降水事件,研究极端干旱和极端降水事件对沙垄不同坡位和坡向短命植物层片生产力的影响。结果表明: 极端干旱使地上净初级生产力和地下净初级生产力分别显著降低48.8%和13.7%,极端降水使地上净初级生产力和地下净初级生产力分别显著增加37.9%和23.2%。地上净初级生产力对极端干旱和极端降水的敏感性(0.26和0.21 g·m^-2·mm^-1)显著强于地下净初级生产力的敏感性(0.02和0.03 g·m^-2·mm^-1)。沙垄东坡地上净初级生产力(24.22 g·m^-2)和地下净初级生产力(5.77 g·m^-2)与西坡相比显著增大29.7%和71.7%,而地上净初级生产力和地下净初级生产力对降水变化的敏感性在不同坡位和坡向之间差异不显著。     

Changes in snowmelt date and summer precipitation affect the flowering phenology of Erythronium grandiflorum (glacier lily; Liliaceae)

? Premise of the study: Climate change has affected species worldwide, including alterations in phenology, migration patterns, distribution, and survival. Because Erythronium grandiflorum is an early-season bloomer, alterations in its phenology may have serious implications for many North American Rocky Mountain communities, including changes in resource availability for pollinators and herbivores. ? Methods: We investigated whether changes in the snowmelt date, summer temperature, and summer precipitation have altered the timing and abundance of flowering in E. grandiflorum by collecting long-term data on floral abundance from 1975-2008 in a series of 2 × 2 m plots at the Rocky Mountain Biological Laboratory (RMBL) in Gothic, Colorado in the United States. ? Key results: Snowmelt date and mean summer temperature were negatively correlated. Over the 30-yr study, the snowmelt date advanced by 4.14 d/decade, and mean summer temperature increased by 0.38°C/decade. Summer precipitation was variable, showing no change. The first, peak, and last flowering dates of E. grandiflorum advanced an average of 3.2 d/decade. Furthermore, earlier snowmelt and greater summer precipitation in the previous year led to earlier flowering in E. grandiflorum. There was no change in flowering abundance in this species, indicating it may be controlled by a complex set of abiotic and biotic variables. ? Conclusions: Our study indicates that snowmelt is arriving earlier at the RMBL, which has caused earlier flowering in E. grandiflorum. Because alterations in phenology can disrupt important ecological interactions, information on potential phenological shifts in species that interact with E. grandiflorum is essential in determining the net effect of climate-driven alterations in phenology.     

Volume-based pollen size analysis: an advanced method to assess somatic and gametophytic ploidy in flowering plants

Pollen size is often used as a biological parameter to estimate the ploidy and viability of mature pollen grains. In general, pollen size quantification is performed one- or two-dimensionally using image-based diameter measurements. As these approaches are elaborate and time consuming, alternative approaches that enable a quick, reliable analysis of pollen size are highly relevant for plant research. In this study, we present the volume-based particle size analysis technique as an alternative method to characterize mature pollen. Based on a comparative assay using different plant species (including tomato, oilseed rape, kiwifruit, clover, among others), we found that volume-based pollen size measurements are not biased by the pollen shape or position and substantially reduce non-biological variation, allowing a more accurate determination of the actual pollen size. As such, volume-based particle size techniques have a strong discriminative power in detecting pollen size differences caused by alterations in the gametophytic ploidy level and therefore allow for a quick and reliable estimation of the somatic ploidy level. Based on observations in Arabidopsis thaliana gametophytic mutants and differentially reproducing Boechera polyantha lines, we additionally found that volume-based pollen size analysis provides quantitative and qualitative data about alterations in male sporogenesis, including aneuploid and diploid gamete formation. Volume-based pollen size analysis therefore not only provides a quick and easy methodology to determine the somatic ploidy level of flowering plants, but can also be used to determine the mode of reproduction and to quantify the level of diplogamete formation.     

Phenotypic responses of wild barley to experimentally imposed water stress

Responses to water stress within a population of wild barley from Tabigha, Israel, were examined. The population's distribution spans two soil types: Terra Rossa (TR) and Basalt (B). Seeds were collected from plants along a 100 m transect; 24 genotypes were sampled from TR and 28 from B. Due to different soil water-holding capacities, plants growing on TR naturally experience more intense drought than plants growing on B. In a glasshouse experiment, water was withheld from plants for two periods (10 d and 14 d) after flag leaf emergence. A total of 15 agronomic, morphological, developmental, and fertility related traits were examined by analysis of variance (ANOVA). Ten of these traits were significantly affected by the treatment. A high degree of phenotypic variation was found in the population with significant genotypextreatment and soil typextreatment interactions. Principal component analysis (PCA) was performed using combined control and stress treatment data sets. The first three principal components (pc) explained 88.8% of the variation existing in the population with pc1 (47.9%) comprising yield-related and morphological traits, pc2 (22.9%) developmental characteristics and pc3 (18.0%) fertility-related traits. The relative performance of individual genotypes was determined and water stress tolerant genotypes identified. TR genotypes were significantly less affected by the imposed water stress than B genotypes. Moreover, TR genotypes showed accelerated development under water deficit conditions. Data indicate that specific genotypes demonstrating differential responses may be useful for comparative physiological studies, and that TR genotypes exhibiting yield stability may have value for breeding barley better adapted to drought.     

31种木本植物开花物候与系统发育的关系

植物物候通常被认为是由环境因素,如降水、温度和日照长度所决定,然而环境因素是否是物候唯一的决定因素仍然存在很大争议。谱系结构表征了植物在进化上的顺序,该发育时序是否对物候产生影响,当前仍然未知。在调查2016年春季新疆乌鲁木齐市最常见的31种木本植物的初始开花时间、败花时间和开花持续时间的基础上,通过分析植物开花物候的分布特征、开花物候在乔灌木间的差别、以及植物谱系距离与开花物候距离间的关系,试图揭示植物的开花物候和物种谱系(进化)顺序间的关系。结果表明:(1)新疆乌鲁木齐市31种木本植物的初始开花时间为4月18日±9d、败花时间为5月5日±12d、开花持续时间为(16±8)d;(2)乔木的初始开花时间和败花时间的标准差分别均低于灌木,乔木开花物候相对灌木更稳定;(3)乔木的初始开花和败花时间均显著早于灌木(P0.05),但开花持续时间在两者间未有显著性差异(P0.05);(3)31种木本植物间的初始开花时间距离、败花时间距离和开花持续时间距离均与物种谱系距离存在显著线性回归关系(P0.05)。综上可知:乔灌木在垂直空间上的分化使得木本植物的开花物候在植物生活型间存在不同。对植物的开花物候,除已被证明的降水、温度和日照长度等环境因素的影响外,物种进化顺序也可能造成了它在植物种间、时间和空间上的变异。     

1980—2005年中国东北木本植物物候特征及其对气温的响应

基于25年物候观测数据,分析了东北地区木本植物物候的时空变化特征,阐述了近25年来东北地区气候变暖对植物物候的影响,结果表明:1980—2005年,东北地区木本植物展叶初期主要呈提前趋势,提前幅度为0.23d·a-1,枯黄初期主要表现为推后趋势,平均推后0.19d·a-1,生长季延长,平均延长幅度为0.30d·a-1;东北地区广泛分布的5种木本植物(旱柳、杏树、小叶杨、榆树和紫丁香)物候在地理空间上存在显著差异:纬度平均每增加1°,展叶初期推后3d,枯黄初期提前1.35d,生长季长度缩短4.41d;东北地区植物展叶初期与2、3、4月的气温显著负相关(P0.05),其中,4月气温对植物展叶提前的影响最大,展叶提前趋势平均为2.35d·℃-1;2月气温的影响最小,平均趋势为1.18d·℃-1。     

植物物候对气候变化的响应

植物物候的变化可以直观地反映某些气候变化,尤其是气候变暖.植物生长节律的变化引起植物与环境关系的改变.生态系统的物质循环（如水和碳的循环）等过程将随物候而改变.不同种类植物物候对气候变化的响应的差异,会使植物间和动植物间的竞争与依赖关系也发生深刻的变化.目前欧洲、美洲、亚洲等许多地区均有关于春季植物物候提前,秋季物候推迟,使植物的生长季延长,从而提示气候变暖的趋势.植物物候的模拟模型构成生态系统生产力模型的重要部分.     

Experimental evaluation of seed limitation in alpine snowbed plants

Background

The distribution and abundance of plants is controlled by the availability of seeds and of sites suitable for establishment. The relative importance of these two constraints is still contentious and possibly varies among species and ecosystems. In alpine landscapes, the role of seed limitation has traditionally been neglected, and the role of abiotic gradients emphasized.

Methodology/Principal Findings

We evaluated the importance of seed limitation for the incidence of four alpine snowbed species (Achillea atrata L., Achillea clusiana Tausch, Arabis caerulea L., Gnaphalium hoppeanum W. D. J. Koch) in local plant communities by comparing seedling emergence, seedling, juvenile and adult survival, juvenile and adult growth, flowering frequency as well as population growth rates λ of experimental plants transplanted into snowbed patches which were either occupied or unoccupied by the focal species. In addition, we accounted for possible effects of competition or facilitation on these rates by including a measure of neighbourhood biomass into the analysis. We found that only A. caerulea had significantly lower seedling and adult survival as well as a lower population growth rate in unoccupied sites whereas the vital rates of the other three species did not differ among occupied and unoccupied sites. By contrast, all species were sensitive to competitive effects of the surrounding vegetation in terms of at least one of the studied rates.

Conclusions/Significance

We conclude that seed and site limitation jointly determine the species composition of these snowbed plant communities and that constraining site factors include both abiotic conditions and biotic interactions. The traditional focus on abiotic gradients for explaining alpine plant distribution hence appears lopsided. The influence of seed limitation on the current distribution of these plants casts doubt on their ability to readily track shifting habitats under climate change unless seed production is considerably enhanced under a warmer climate.     

Apomixis in flowering plants: an overview

Apomixis is a common feature of perennial plants, which occurs in ca. 60% of the British flora, but has been largely ignored by reproductive theoreticians. Successful individuals may cover huge areas, and live to great ages, favoured by 'symmetrical' selection. Apomixis is favoured by colonizing modes, for instance post-glacially. Despite its theoretical advantages, apomixis usually coexists with sexuality, suggesting 'hidden' disadvantages. Agamospermy (apomixis by seed) is relatively uncommon, but gains from the attributes of the seed. It pays agamospermy genes, which discourage recombination, to form co-adapted linkage groups, so that they become targets for disadvantageous recessive mutant accumulation. Consequently, agamospermy genes cannot succeed in diploids and agamosperms are hybrid and highly heterotic. Agamospermous endosperm may suffer from genomic imbalance, so that nutritious ovules, which can support embryos without endosperm, may be preadapted for agamospermy. When primary endosperm nucleus fertilization ('pseudogamy') continues as a requirement for many aposporous agamosperms, selfing sex becomes preadaptive and archesporial sex remains an option. Apomictic populations can be quite variable although apomictic families are much less variable than sexuals. Only in some diplosporous species does sex disappear completely, and in those species some release of variability may persist through somatic recombination. The search for an agamospermy gene suitable for genetic modification should target fertile sexuals with a single localized agamospermy (A) gene, which therefore lack a genetic load. The A gene should coexist alongside sexuality, so that it would be easy to select seedlings of sexual and asexual origins. Plants with sporophytic agamospermy provide all these attributes.     

Resource-related variables drive individual variation in flowering phenology and mediate population-level flowering responses to climate in an asynchronously reproducing palm

Many tropical plant species show wide intra-population variation in reproductive timing, resulting in the protracted presence of flowering and fruiting individuals. Various eco-evolutionary drivers have been proposed as ultimate causes for asynchronous phenology, yet little is known about the proximate factors that control reproductive onset among individuals or that influence the proportion of trees producing new inflorescences within a population. We employed a nine-year phenological record from 178 individuals of the hyperdominant, asynchronously flowering canopy palm, Oenocarpus bataua (Arecaceae)¸ to assess whether resource-related variables influence individual- and population-level flowering phenology. Among individuals, access to sunlight increased rates of inflorescence production, while the presence of resource sinks related to current investment in reproduction—developing infructescences—reduced the probability of producing new inflorescences. At the population level, climate anomalies induced by El Niño Southern Oscillation (ENSO) affected the proportion of the population producing inflorescences through time. Moreover, the effects of ENSO anomalies on flowering patterns depended on the prevalence of developing infructescences in the population, with stronger effects in periods of low developing-infructescence frequency. Taken together, these results suggest that resource-related variables can drive phenological differences among individuals and mediate population-level responses to larger-scale variables, such as climate anomalies. Consequently, a greater focus on the role of resource levels as endogenous cues for reproduction might help explain the frequent aseasonal phenological patterns observed among tropical plants, particularly those showing high intra-population asynchrony.