Alpine Plant Research Articles

Multi-Omics Analysis Reveals the Molecular Mechanisms of the Glycolysis and TCA Cycle Pathways in Rhododendron chrysanthum Pall. under UV-B Stress

UV-B radiation is becoming a bigger threat to plants as a result of the ozone layer’s depletion. As an alpine plant, Rhododendron chrysanthum Pall. (R. chrysanthum) may grow regularly under UV-B radiation throughout its lengthy acclimatization period, although the mechanism of acclimatization is still poorly understood. The current investigation uncovered a number of adaptation strategies that R. chrysanthum has developed in reaction to UV-B rays. UV-B radiation impeded photosynthesis and damaged the photosystem, according to OJIP testing. Through transcriptomics and proteomics analyses, this study found that the differential proteins and differential genes of R. chrysanthum were significantly enriched in glycolysis and tricarboxylic acid (TCA) cycle pathways after UV-B treatment. The metabolomics results showed that a total of eight differential metabolites were detected in the glycolytic and TCA cycle pathways, and the changes in the expression of these metabolites reflected the final outcome of gene regulation in the glycolytic and TCA cycle pathways. The combined experimental results demonstrated that R. chrysanthum’s photosynthetic system was impacted by UV-B stress and, concurrently, the plant activated an adaptation mechanism in response to the stress. To maintain its energy supply for growth, R. chrysanthum adapts to UV-B stress by adjusting the expression of the relevant proteins, genes, and metabolites in the glycolytic and TCA cycling pathways. This study provides a new perspective for understanding the changes in the carbon metabolism of R. chrysanthum under UV-B stress and its mechanisms for UV-B resistance, and provides an important theoretical basis for the study of enhancing plant resistance to stress.

The complete plastome of Saxifraga giraldiana Engler and its phylogenetic analysis

Saxifraga giraldiana Engler is a common subalpine and alpine plant belonging to Saxifragaceae. However, the genetic diversity of this species has remained to be explored. In this study, we have assembled and characterized the complete chloroplast genome of S. giraldiana, filling this knowledge gap and uncovering its genetic composition. The chloroplast genome is 147,267 bp long and contains 131 genes, including 85 protein-coding genes, 38 tRNA genes, and eight rRNA genes. Furthermore, we have performed a phylogenetic analysis of 19 representative species within Saxifraga. As a result, we have found that S. giraldiana, together with S. implicans and S. stellariifolia, forms a monophyletic group. These findings have implications for the conservation and utilization of S. giraldiana.

Response of alpine vegetation function to climate change in the Tibetan Plateau: A perspective from solar-induced chlorophyll fluorescence

Vegetation change in the Tibetan Plateau (TP) is a crucial indicator of climate change in alpine regions. Previous studies have reported an overall greening trend in the vegetation structure across the TP, especially in its northeastern part, in response to a warming climate. However, variations in the vegetation function and the possible drivers remain poorly understood. Considering the optimal temperature for plants in TP is usually higher than the current temperature, our hypothesis is the function and structure of alpine vegetation have changed synchronously over past few decades. To test this hypothesis, we analyzed satellite-observed solar-induced chlorophyll fluorescence (SIF) and leaf area index (LAI) in the Yellow River source (YRS) region in the northeastern TP to quantify the long-term trends in vegetation functional and structural states, respectively. The results suggest that from 1982 to 2018, SIF increased significantly in 77.71 % of the YRS area, resulting in a significant upward trend of 0.52 × 10−3 mW m−2 nm−1 sr−1 yr−1 (p < 0.001) for the regional-mean SIF. This represents a 16.1 % increase in SIF, which is close in magnitude to the increase in LAI over the same period. The synchronous changes between vegetation function and structure suggest that improved greenness corresponds to a similar level of change in carbon uptake across YRS. Additionally, we used a multiple regression approach to quantify the contribution of climatic factors to SIF changes in YRS. Our analyses show that the increases in SIF were primarily driven by rising temperatures. Spatially, temperature dominated SIF changes in most parts of YRS, except for certain dry parts in the northern and western YRS, where precipitation had a greater impact. Our results are crucial for a comprehensive understanding of climate regulations on vegetation structure and function in high-elevation regions.

Fine‐scale alpine plant community assembly: Relative roles of environmental sorting, dispersal processes and species interactions

Abstract Besides environmental sorting, other processes like biotic interactions and dispersal limitation are vital for the assembly of plant communities in high mountains and their re‐assembly under changing climatic conditions. Nevertheless, studies that compare the impact of these factors on plant community assembly above the tree line are largely lacking so far. We analysed occurrence changes in vascular plant communities of 492 permanent 1‐m2 plots in the alpine‐nival ecotone of Mt. Schrankogel, Austrian Alps by comparing resurvey data from 2014 with data from the initial survey in 1994. We combined these data with species inventories from 899 additional plots sampled in 2021 and 2022 across a larger landscape above the tree line covering an elevational range of 1700 m, which we used for fine‐scale habitat suitability modelling. We assessed the relative effects of projected habitat suitability, propagule pressure from surrounding populations and biomass density of neighbours on 1532 colonization and 372 extirpation events of 31 species observed on the permanent plots. We found that all three factors are significantly related to both colonisations and extirpations, with habitat suitability having the strongest, propagule pressure a slightly weaker, and vegetation density the weakest effect. Colonisations can be better explained by the three process proxies than extirpations. Our results indicate a crucial role of dispersal limitation besides the predominant effect of environmental filtering on the (re‐)assembly of the alpine‐nival plant community, while competitive/facilitative effects between plants tend to play a minor role. The strong imprint of nearby source plant populations on colonization/extirpation events suggests that recent plant migrations predominantly occur in small steps. This implies that while the topographically complex alpine terrain offers climatic microrefugia for plants, it may also pose potential barriers, hindering species from following their suitable climatic niches upwards. Synthesis: Besides filtering by environmental conditions dispersal limitation had a strong effect on the observed changes in a local alpine plant community over two decades. Limited dispersal capacities of plant species may counteract the ability of isolated cold areas to effectively shelter high alpine plants from the effects of climate warming.

Transcriptome analysis unveiled the genetic basis of rapid seed germination strategies in alpine plant Rheum pumilum

Rheum pumilum stands as both a quintessential alpine plant and a significant traditional Chinese and Tibetan medicinal herb. Unraveling the molecular intricacies of seed germination in Rh. pumilum not only unveils the genetic foundations of plant seed germination strategies in high-altitude environments but also offers insights for cultivating Rh. pumilum medicinal materials. Employing transcriptome sequencing and the Weighted Gene Co-expression Network Analysis, this study delved into the shifts in gene expression levels across various stages of seed germination in Rh. pumilum. The process of seed germination in Rh. pumilum entails a cascade of complex physiological events. Six hormones (ABA, IAA, ETH, GA, BR, CK) emerged as pivotal players in seeds breaking in shells and the facilitation of rapid seed germination in Rh. pumilum. Fourteen transcription factor families (LOB, GRAS, B3, bHLH, bZIP, EIL, MYB, MYB related, NAC, TCP, WRKY, HSF, PLATZ, and SBP) along with four key genes (E2.4.1.13, EIN3, BZR, and BIN2) were identified that may be associated with both biotic and abiotic environmental stress. The ETR, ACACA and ATPeV0C genes were linked with energy accumulation during the initial stages of seed germination, CYP707A may play an important role in breaking seed dormancy, while the BRI1 gene may be correlated with swift seed germination. Additionally, several unidentified genes were recognized to play key roles in seed germination of Rh. pumilum, warranting further investigation. Moreover, Rh. pumilum demonstrates full activation of crucial physiological functions such as energy metabolism, signal transduction, and responses to biological and abiotic stresses during the seed breaking in shells. This study provides molecular evidence elucidating the swift seed germination strategies adopted by alpine plants to thrive in high-altitude environments. Furthermore, it serves as a foundational reference for enhancing seed germination rates and breeding practices to promote the sustainable development of Rh. pumilum medicinal materials.

Fine-Scale Lithogeochemical Features Influence Plant Distribution Patterns in Alpine Grasslands in the Western Alps of Italy.

Bedrock geology is crucial in structuring alpine plant communities. Old studies mainly focused on the compositional differences between alpine plant communities on carbonate rocks and crystalline rocks, i.e., calcareous vs. siliceous vegetation. Increasing attention is being paid to bedrock types other than calcareous or siliceous ones, viz. those which have intermediate geochemical characteristics between pure calcareous and pure siliceous ones. Among these types of 'intermediate' bedrocks, calc-schists and serpentines are generally characterized by vegetation comprised of a mixture of basiphilous and acidophilous species. We selected several sites in alpine grasslands in the Western Italian Alps, on calc-schist and serpentine bedrocks, located at 2500 ± 100 m above sea level. X-ray fluorescence quantification of major and trace elements, combined with stereomicroscopic examination of bedrock samples with a petrographic approach, revealed a much broader range of bedrock types than recognized by inspection of geological maps. The vegetation investigated in our study was mostly composed of a set of species found more or less frequently in alpine silicicolous or calcicolous plant communities of the Alps and other European mountains. The carbonate content in the bedrock was one of the main drivers of variation in grassland vegetation, not necessarily related to soil pH. There were no distinctive species uniquely characterizing grassland vegetation on serpentines or calc-schists.

Growth dynamics and climate sensitivities in alpine cushion plants: insights from Silene acaulis in the Swiss Alps

AbstractUnderstanding alpine plants’ growth dynamics and responses to warming is essential for predicting climate change impacts on mountain ecosystems. Here, we examine growth determinants in the alpine cushion plant Silene acaulis in the Swiss Alps, exploring ontogeny, elevation, and climate influences. We collected 40 Silene individuals and 159 individuals from 38 co-occurring alpine species across 2200–3130 m elevations in the Swiss Alps, analyzing age and growth histories through annual growth rings. While comparing growth rates, we found that Silene was relatively slow-growing. However, Silene exhibited a dual growth strategy, initially rapid and then slowing after ~ 20 years, challenging perceptions of its longevity. Similar ontogenetic trends were observed in other alpine species, albeit with variations based on species and elevation. The consistent unimodal growth-elevation pattern in Silene and other alpine plants, peaking at ~ 2400 m, underscores shared environmental constraints on alpine plant growth. Additionally, cross-dating growth ring series and correlating with daily climate data enabled the precise assessment of warming impacts on growth. Silene’s growth is influenced by year-to-year climate variability, with warming-induced moisture stress and overheating during spring and summer adversely affecting its growth. Despite being low-statured, Silene is not completely decoupled from atmospheric influences. The heat-trapping function of Silene, effective in mature and well-formed cushions, makes it susceptible to adverse effects as temperatures rise. This sensitivity raises concerns about the potential dieback of Silene cushions, as witnessed during recent heatwaves, and emphasizes the broader ecological implications for alpine ecosystems, given Silene’s role as a crucial nurse plant.

Snow drifts as a driver of alpine plant productivity as observed from weekly multispectral drone imagery

Snow drifts as a driver of alpine plant productivity as observed from weekly multispectral drone imagery

Anticoccidial activity of the secondary metabolites in alpine plants frequently ingested by wild Japanese rock ptarmigans

The Japanese rock ptarmigan (Lagopus muta japonica) is an herbivorous species of partridges that inhabits only alpine zones. Alpine plants are their main source of food. These alpine plants contain toxic compounds to deter herbivores from consuming them. A previous analysis of the alpine plants frequently consumed by Japanese rock ptarmigans revealed the presence of a unique mixture of secondary metabolites and a novel compound. Additionally, wild Japanese rock ptarmigans are often infected by two species of Eimeria parasites. When these parasites were experimentally administered to Svalbard rock ptarmigans (Lagopus muta hyperborean), which do not feed on alpine plants, the birds exhibited symptoms, such as diarrhea and depression, and in some cases, they died. Although little is known about the pathogenesis of these parasites in wild Japanese rock ptarmigans, it was hypothesized that compounds found in alpine plants, their main food source, may reduce the pathogenicity of Eimeria parasites. In the present study, we evaluated the anticoccidial activity of the compounds derived from alpine plants in vitro using Eimeria tenella, which infects chickens belonging to the same pheasant family, as an experimental model. Twenty-seven natural components were extracted from eight alpine plants. The natural components were added to E. tenella sporozoites and incubated for 24 h to evaluate their direct effect. Additionally, Madin-Darby bovine kidney cells were incubated with sporozoites and natural components for 24 h to evaluate the inhibitory effect of the components on sporozoite cell invasion. Six compounds from four alpine plants decreased sporozoite viability by up to 88.3%, and two compounds inhibited sporozoite invasion into the cells. Although further studies are needed to evaluate the effects of these components against Eimeria infections in vivo, our findings suggest that these alpine plants may reduce the degree of infection by decreasing the number of sporozoites in the intestinal tract.

Temperature-induced germination pulses above the alpine tree line

ABSTRACT Subject to a long research tradition, the tree line is considered an important biogeographic indicator of climate changes and associated range shifts. Realized tree line positions and the potential tree line isotherm are, however, rarely in equilibrium because trees are unable to track rapid temperature variations. Often ignored in tree line research, this dilemma constrains the suitability of tree line trees for understanding alpine vegetation responses to anthropogenic warming. Here, we present combined dendrochronological and wood anatomical assessments of 1,351 seedlings and saplings from three subalpine forest species—larch (Larix decidua Mill.), pine (Pinus cembra L.), and spruce (Picea abies)—collected between ~2,200 and 2,600 m.a.s.l. in the Swiss Alps. We found evidence for temperature-induced, pulse-like seedling germination, rather than a continuous, long-term upward movement. Though the species spread across overlapping elevational ranges, larch was found at the highest elevations, followed by spruce and pine. Surprisingly, we found a varying age structure, with no sign of decreasing age toward higher elevations. Spring and summer temperatures promoted germination pulses, but postgermination survival was likely facilitated by species-specific plant traits. Our study demonstrates the importance of seedling and sapling data from above the tree line to understand prevailing vegetation dynamics at cold temperature extremes and also suggests future tree line advancement in the Swiss Alps.

Xiaojiang River Basin Ecological Environmental Quality Spatiotemporal Pattern and Evolutionary Trend Analysis Using GEE from 1990 to 2022

Assessment and monitoring of the quality of the ecological environment in the area is a very important fundamental task in the development of ecological civilization in the Xiaojiang River Basin in Yunnan Province, which serves as a demonstration area for ecological restoration in the upper reaches of the Yangtze River. The Landsat remote sensing images from 1990, 1995, 2000, 2005, 2010, 2014, 2018, and 2022 were chosen, and the four indexes of greenness （NDMVI）, humidity （WET）, dryness （NDBSI）, and heat （LST） were extracted. The remote sensing ecological index （RSEI） was created using the principal component analysis method, then the spatial and temporal patterns and trends of ecological quality in the Xiaojiang River Basin between 1990 and 2022 were examined using the GEE platform, ArcGIS 10.7 platform, and Python platform, combining the analysis methods of geographic information mapping, coefficient of variation, Mann-Kendall trend test, Sen's slope estimation, and Hurst's index. The findings demonstrated that： ① the ecological quality of the study area had more obvious geographic differentiation spatially, and by 2022, the areas with excellent and good ecological quality grades were primarily distributed in the areas with better alpine vegetation cover, and those with poor ecological quality were primarily distributed in the areas of the mudslide ravines with relatively low terrain. On a time scale, the study area's RSEI index increased from 0.41 in 1990 to 0.55 in 2022, with a fluctuating overall trend of ecological quality improvement and an average increase of 0.048（10 a） -1； this progress was directly related to a number of ecological construction initiatives that have been energetically carried out, such as converting farms to forests, preventing mudslides, saving soil and water, managing heavy metal contamination, etc. ② The RSEI was more appropriate for the evaluation of ecological quality in alpine ravine areas because, in comparison to the NDVI index, the NDVMI adopted in this study was more sensitive to vegetation information in topographic undulation areas, especially in shaded areas, and could more accurately and quantitatively describe the vegetation information. ③ The RSEI in the Xiaojiang River Basin had a mean coefficient of variation of 0.202. Overall, its volatility was low, and its high volatility was mostly concentrated in the mudslide gully area along both sides of the Xiaojiang River fracture zone, where the surface was made up of bare rocks and sediment that was easily impacted by the changing of the seasons, the climate, and human activity. ④ The quality of the ecological environment in the region was significantly improving, with the rising area reaching 85.72% of the total area and the declining area accounting for approximately 10.15% of the total area. The future trend of change will be dominated by ongoing improvement and future degradation, accounting for 44.75% and 39.97%, respectively. It is important to pay close attention to areas that could potentially degrade. The findings of this study can serve as a theoretical foundation for additional ecological environmental conservation, management, and sustainable development in the Xiaojiang River Basin.

Golden 2-like Transcription Factors Regulate Photosynthesis under UV-B Stress by Regulating the Calvin Cycle.

UV-B stress can affect plant growth at different levels, and although there is a multitude of evidence confirming the effects of UV-B radiation on plant photosynthesis, there are fewer studies using physiological assays in combination with multi-omics to investigate photosynthesis in alpine plants under stressful environments. Golden 2-like (G2-like/GLK) transcription factors (TFs) are highly conserved during evolution and may be associated with abiotic stress. In this paper, we used Handy-PEA and Imaging-PAM Maxi to detect chlorophyll fluorescence in leaves of Rhododendron chrysanthum Pall. (R. chrysanthum) after UV-B stress, and we also investigated the effect of abscisic acid (ABA) on photosynthesis in plants under stress environments. We used a combination of proteomics, widely targeted metabolomics, and transcriptomics to study the changes of photosynthesis-related substances after UV-B stress. The results showed that UV-B stress was able to impair the donor side of photosystem II (PSII), inhibit electron transfer and weaken photosynthesis, and abscisic acid was able to alleviate the damage caused by UV-B stress to the photosynthetic apparatus. Significant changes in G2-like transcription factors occurred in R. chrysanthum after UV-B stress, and differentially expressed genes localized in the Calvin cycle were strongly correlated with members of the G2-like TF family. Multi-omics assays and physiological measurements together revealed that G2-like TFs can influence photosynthesis in R. chrysanthum under UV-B stress by regulating the Calvin cycle. This paper provides insights into the study of photosynthesis in plants under stress, and is conducive to the adoption of measures to improve photosynthesis in plants under stress to increase yield.

Phenological responses of alpine snowbed communities to advancing snowmelt.

Climate change is leading to advanced snowmelt date in alpine regions. Consequently, alpine plant species and ecosystems experience substantial changes due to prolonged phenological seasons, while the responses, mechanisms and implications remain widely unclear. In this 3-year study, we investigated the effects of advancing snowmelt on the phenology of alpine snowbed species. We related microclimatic drivers to species and ecosystem phenology using in situ monitoring and phenocams. We further used predictive modelling to determine whether early snowmelt sites could be used as sentinels for future conditions. Temperature during the snow-free period primarily influenced flowering phenology, followed by snowmelt timing. Salix herbacea and Gnaphalium supinum showed the most opportunistic phenology, while annual Euphrasia minima struggled to complete its phenology in short growing seasons. Phenological responses varied more between years than sites, indicating potential local long-term adaptations and suggesting these species' potential to track future earlier melting dates. Phenocams captured ecosystem-level phenology (start, peak and end of phenological season) but failed to explain species-level variance. Our findings highlight species-specific responses to advancing snowmelt, with snowbed species responding highly opportunistically to changes in snowmelt timings while following species-specific developmental programs. While species from surrounding grasslands may benefit from extended growing seasons, snowbed species may become outcompeted due to internal-clock-driven, non-opportunistic senescence, despite displaying a high level of phenological plasticity.

Testing the evolutionary potential of an alpine plant: phenotypic plasticity in response to growth temperature outweighs parental environmental effects and other genetic causes of variation.

Phenotypic plasticity and rapid evolution are fundamental processes by which organisms can maintain their function and fitness in the face of environmental changes. Here we quantified the plasticity and evolutionary potential of an alpine herb Wahlenbergia ceracea. Utilizing its mixed-mating system, we generated outcrossed and self-pollinated families that were grown in either cool or warm environments, and that had parents that had also been grown in either cool or warm environments. We then analysed the contribution of environmental and genetic factors to variation in a range of phenotypic traits including phenology, leaf mass per area, photosynthetic function, thermal tolerance, and reproductive fitness. The strongest effect was that of current growth temperature, indicating strong phenotypic plasticity. All traits except thermal tolerance were plastic, whereby warm-grown plants flowered earlier, grew larger, and produced more reproductive stems compared with cool-grown plants. Flowering onset and biomass were heritable and under selection, with early flowering and larger plants having higher relative fitness. There was little evidence for transgenerational plasticity, maternal effects, or genotype×environment interactions. Inbreeding delayed flowering and reduced reproductive fitness and biomass. Overall, we found that W. ceracea has the capacity to respond rapidly to climate warming via plasticity, and the potential for evolutionary change.

The polyphyletic Caucasus-centred Campanula subg. Scapiflorae (Campanulaceae) revisited with a newly circumscribed C. sect. Tridentatae for its core clade.

Campanula L. is among the genera with the highest number of endemics in the Caucasus ecoregion. A group of attractive alpine and subalpine perennial rosette plants with short single-flowered stems centred in the Caucasus has been treated as Campanulasubg.Scapiflorae or at other ranks, with considerably varying circumscription and classification. Molecular phylogenetic analysis of three plastid DNA regions (trnK/matK, petD, rpl16) of a strongly extended sampling, comprising 23 of the 27 commonly accepted taxa (85%) with 330 accessions built on and guided by the results of our previous study of the group, confirmed the polyphyly of C.subg.Scapiflorae in any of its circumscriptions. The core clade of the group comprises exclusively endemics and near-endemics of the Caucasus and is treated here as C.sect.Tridentatae in a revised circumscription. The phylogenetic relationships of the disparate other elements of the Scapiflorae group are outlined.

Short-Term Responses of Alpine Vegetation to the Removal of Dominant versus Sparse Species.

The mass ratio hypothesis posits that ecosystem functions are predominantly influenced by the dominant species. However, it remains unclear whether a species must be abundant to exert functional dominance. We conducted a removal experiment in an alpine grassland near Pudacuo National Park, Yunnan, China, to assess the community and ecosystem impacts of the removed species. We implemented four treatments as follows: exclusive removal of the most abundant species (Blysmus sinocompressus), exclusive removal of a sparse species with high individual biomass (Primula secundiflora), simultaneous removal of both species, and a control with no removals. Results showed that removing B. sinocompressus significantly reduced biomass production, supporting the mass ratio hypothesis, while removal of P. secundiflora had negligible effects. B. sinocompressus removal positively impacted community metrics like coverage, species evenness, and the Shannon diversity index, but not species richness, likely due to its spatial dominance. Conversely, P. secundiflora removal had minimal community impact, probably due to its limited influence on nearby species. This study underscores the proportionate roles of the dominant species in alpine grasslands, emphasizing that their community and ecosystem impacts are proportional to their abundance.

Environmental factors shaping alpine plant adaptations

Alpine ecosystems are distinct and demanding settings that are defined by severe climatic extremes, such as frigid temperatures, intense radiation, and restricted nutrition supply. The adaptations of alpine plants are finely tuned responses to the unique environmental challenges they face. By delving into the intricate interplay between temperature, UV radiation, soil conditions, and precipitation patterns, are valuable insights into the resilience of alpine ecosystems. Such knowledge is not only crucial for advancing ecological understanding but also for informing conservation and management strategies in the era of global environmental change. We have covered all the abiotic factors affecting plant growth and development in alpine and subalpine area in this mini review. It will be beneficial for researchers, industrialists, ecologists and agriculturists.

Spatial patterns and variations in leaf traits of alpine plants on the interior Tibetan Plateau

Spatial patterns and environmental gradients in plant traits can elucidate the relationships between plant characteristics, environmental factors, and ecosystem functions and can be used to develop the next generation of vegetation models for predicting vegetation distribution and functions. Studies of Plant traits across spatial scales have advanced significantly worldwide, including on the Tibetan Plateau (TP), the highest and largest plateau globally. Previous research on plant traits on the TP has been limited to specific areas, leaving the overall trend across the plateau unclear. In this study, we examined seven leaf traits—leaf thickness (LT), leaf dry matter content (LDMC), leaf area (LA), specific leaf area (SLA), and leaf carbon (LCC), nitrogen (LNC), and phosphorus (LPC) contents—in 1314 individuals at 237 sites within the alpine vegetation region above 3000 m on the TP to determine their distribution patterns across three spatial dimensions and their relationships with environmental factors. Our results indicated that, compared with Chinese plants, TP plants exhibited significantly greater LT, LDMC, and LNC but lower LA, LCC, and LPC. Except for SLA and LCC, most leaf traits showed significant spatial variability. Climate and soil explained 29.23 % of the total variations in traits. Radiation had the most pronounced effect on LDMC, LA, LNC, and LPC, whereas soil pH primarily constrained LT and LCC. Precipitation influenced most traits both directly and indirectly, with substantial overall effects. Temperature and soil nutrient had minimal influence on the regulation of leaf traits across most of the TP. Our findings suggest that alpine plants typically employ a conservative strategy to cope with the harsh conditions on the TP, but this strategy varies at broader spatial scales and longer environmental gradients, particularly in extreme environments such as the highest alpine regions.

Ecosystem engineers can regulate resource allocation strategies in associated plant species.

Balancing the biomass requirements of different functions for the purpose of population reproduction and persistence can be challenging for alpine plants due to extreme environmental stresses from both above- and below-ground sources. The presence of ecosystem engineers in alpine ecosystems effectively alleviates microenvironmental stresses, hence promoting the survival and growth of other less stress-tolerant species. However, the influence of ecosystem engineers on plant resource allocation strategies remains highly unexplored. In this study, we compared resource allocation strategies, including biomass accumulation, reproductive effort (RE), root fraction (RF), as well as relationships between different functions, among four alpine plant species belonging to Gentianaceae across bare ground, tussock grass-, cushion-, and shrub-engineered microhabitats. Shrub-engineered microhabitats exerted the strongest effects on regulating plant resource allocation patterns, followed by tussock grass- and cushion-engineered microhabitats. Additionally, apart from microhabitats, population background and plant life history also significantly influenced resource allocation strategies. Generally, plants established within engineered microhabitats exhibited higher biomass accumulation, as well as increased flower, leaf and stem production. Furthermore, individuals within engineered microhabitats commonly displayed lower RF, indicating a greater allocation of resources to above-ground functions while reducing allocation to root development. RE of annual plants was significantly higher than that of perennial plants. However, individuals of annual plants within engineered microhabitats showed lower RE compared to their counterparts in bare ground habitats; whereas perennial species demonstrated similar RE between microhabitat types. Moreover, RE was generally independent of plant size in bare-ground habitats but exhibited size-dependency in certain populations for some species within specific engineered microhabitat types. However, size-dependency did exist for absolute reproductive and root biomass allocation in most of the cases examined here. No trade-offs were observed between flower mass and flower number, nor between leaf mass and leaf number. The capacity of ecosystem engineers to regulate resource allocation strategies in associated plants was confirmed. However, the resource allocation patterns resulted synergistically from the ecosystem engineering effects, population environmental backgrounds, and plant life history strategies. In general, such regulations can improve individual survival and reproductive potential, potentially promoting population persistence in challenging alpine environments.

Climate refugia along Lake Superior’s shores: disjunct arctic–alpine plants rely on cool shoreline temperatures but are restricted to highly exposed habitat under climate warming

Abstract Climate refugia can serve as a remnant habitat or stepping stones for species dispersal under climate warming. The largest freshwater lake by surface area, Lake Superior, USA and Canada, serves as a model system for understanding cooling-mediated local refugia, as its cool water temperatures and wave action have maintained shoreline habitats suitable for southern disjunct populations of arctic–alpine plants since deglaciation. Here, we seek to explain spatial patterns and environmental drivers of arctic–alpine plant refugia along Lake Superior’s shores, and assess future risk to refugia under moderate (+3.5 °C) and warmest (+5.7 °C) climate warming scenarios. First, we examined how the interactive effects of summer surface water temperatures and wind affected onshore temperatures, resulting in areas of cooler refugia. Second, we developed an ecological niche model for the presence of disjunct arctic–alpine refugia (pooling 1253 occurrences from 58 species) along the lake’s shoreline. Third, we fit species distribution models for 20 of the most common arctic–alpine disjunct species and predicted presence to identify refugia hotspots. Finally, we used the two climate warming scenarios to predict changes in the presence of refugia and disjunct hotspots. Bedrock type, elevation above water, inland distance, July land surface temperature from MODIS/Terra satellite and near-shore depth of water were the best predictors of disjunct occurrences. Overall, we predicted 2236 km of the shoreline (51%) as disjunct refugia habitat for at least one species under current conditions, but this was reduced to 20% and 7% with moderate (894 km) and warmest (313 km) climate change projections.