Four ways to define the growing season.

What is addressed as growing season in terrestrial ecosystems is one of the main determinants of annual plant biomass production globally. However, there is no well-defined concept behind. Here, we show different facets of what might be termed growing season, each with a distinct meaning: (1) the time period during which a plant or a part of it actually grows and produces new tissue, irrespective of net carbon gain (growing season sensu stricto). (2) The period defined by developmental, that is, phenological markers (phenological season). (3) The period during which vegetation as a whole achieves its annual net primary production (NPP) or a net ecosystem production (NEP), expressed as net carbon gain (productive season) and (4) the period during which plants could potentially grow based on meteorological criteria (meteorological season). We hypothesize that the duration of such a 'window of opportunity' is a strong predictor for NPP at a global scale, especially for forests. These different definitions have implications for the understanding and modelling of plant growth and biomass production. The common view that variation in phenology is a proxy for variation in productivity is misleading, often resulting in unfounded statements on potential consequences of climatic warming such as carbon sequestration.

Growth of alpine grassland will start and stop earlier under climate warming.

Alpine plants have evolved a tight seasonal cycle of growth and senescence to cope with a short growing season. The potential growing season length (GSL) is increasing because of climate warming, possibly prolonging plant growth above- and belowground. We tested whether growth dynamics in typical alpine grassland are altered when the natural GSL (2-3 months) is experimentally advanced and thus, prolonged by 2-4 months. Additional summer months did not extend the growing period, as canopy browning started 34-41 days after the start of the season, even when GSL was more than doubled. Less than 10% of roots were produced during the added months, suggesting that root growth was as conservative as leaf growth. Few species showed a weak second greening under prolonged GSL, but not the dominant sedge. A longer growing season under future climate may therefore not extend growth in this widespread alpine community, but will foster species that follow a less strict phenology.

Snowmaking in a warmer climate: an in-depth analysis of future water demands for the ski resort Andermatt-Sedrun-Disentis (Switzerland) in the twenty-first century.

Rising air temperatures threaten the snow reliability of ski resorts. Most resorts rely on technical snowmaking to compensate lacking natural snow. But increased water consumption for snowmaking may cause conflicts with other sectors' water uses such as hydropower production or the hotel industry. We assessed the future snow reliability (likelihood of a continuous 100-day skiing season and of operable Christmas holidays) of the Swiss resort Andermatt-Sedrun-Disentis throughout the twenty-first century, where 65% of the area is currently equipped for snowmaking. Our projections are based on the most recent climate change scenarios for Switzerland (CH2018) and the model SkiSim 2.0 including a snowmaking module. Unabated greenhouse gas emissions (scenario RCP8.5) will cause a lack of natural snow at areas below 1800-2000 m asl by the mid-twenty-first century. Initially, this can be fully compensated by snowmaking, but by the end of the century, the results become more nuanced. While snowmaking can provide a continuous 100-day season throughout the twenty-first century, the economically important Christmas holidays are increasingly at risk under the high-emission scenario in the late twenty-first century. The overall high snow reliability of the resort comes at the cost of an increased water demand. The total water consumption of the resort will rise by 79% by the end of the century (2070-2099 compared to 1981-2010; scenario RCP8.5), implying that new water sources will have to be exploited. Future water management plans at the catchment level, embracing the stakeholders, could help to solve future claims for water in the region.

Biomass allocation and seasonal non-structural carbohydrate dynamics do not explain the success of tall forbs in short alpine grassland.

The majority of alpine plants are of small stature. Through their small size alpine plants are decoupled from the free atmospheric circulation and accumulate solar heat. However, a few alpine species do not follow that "rule" and protrude with their aboveground structures from the microclimatic shelter of the main canopy boundary layer. We aim at explaining the phenomenon of being tall by exploring the biomass production and carbon relations of four pairs of small and tall phylogenetically related taxa in alpine grassland. We compared species and stature-specific biomass allocation, shifts in non-structural carbohydrate (NSC) concentrations in different tissues throughout the season, and we used 13C labels to track carbon transfer from leaves to belowground structures. Small and tall herbs did not differ in their above- to belowground biomass allocation. The NSC composition (starch, fructan, simple sugars) and allocation did not show a stature-specific pattern, except for higher concentrations of simple sugars in tall species during their extended shoot growth. In relative terms, tall species had higher NSC pools in rhizomes, whereas small species had higher NSC pools in roots. Our findings do not place tall alpine forbs in an exceptional category in terms of biomass allocation and carbohydrate storage. The tall versus small stature of the examined herbs does not seem to be associated with specific adjustments in carbon relations. 13C pulse labelling revealed early C autonomy in young, unfolding leaves of the tall species, which are thus independent of the carbon reserves in the massive belowground organs.

Halving sunlight reveals no carbon limitation of aboveground biomass production in alpine grassland.

In temperate alpine environments, the short growing season, low temperature and a slow nutrient cycle may restrict plant growth more than carbon (C) assimilation does. To test whether C is a limiting resource, we applied a shade gradient from ambient light to 44% (maximum shade) of incident photon flux density (PFD) in late successional, Carex curvula-dominated alpine grassland at 2,580 m elevation in the Swiss central Alps for 3 years (2014-2016). Total aboveground biomass did not significantly decrease under reduced PFD, with a confidence interval ranging from +4% to -15% biomass in maximum shade. Belowground biomass, of which more than 80% were fine roots, was significantly reduced by a mean of 17.9 ± 4.6% (±SE), corresponding to 228 g/m2 , in maximum shade in 2015 and 2016. This suggests reduced investments into water and nutrient acquisition according to the functional equilibrium concept. Specific leaf area (SLA) and maximum leaf length of the most abundant species increased with decreasing PFD. Foliar concentration of nonstructural carbohydrates (NSC) was reduced by 12.5 ± 4.3% under maximum shade (mean of eight tested species), while NSC concentration of belowground storage organs were unchanged in the four most abundant forbs. Furthermore, maximum shade lowered foliar δ13 C by 1.56 ± 0.35‰ and increased foliar nitrogen concentrations per unit dry mass by 18.8 ± 4.1% across six species in 2015. However, based on unit leaf area, N concentrations were lower in shade (effect of higher SLA). Thus, while we found typical morphological and physiological plant responses to lower light, shading did not considerably affect seasonal aboveground biomass production of this alpine plant community within a broad range of PFD. This suggests that C is not a growth-limiting resource, matching the unresponsiveness to in situ CO2 enrichment previously reported for this type of grassland.

Legacy Effects of Climate Extremes in Alpine Grassland.

Climate change is particularly apparent in many mountainous regions, with warming rates of more than twice the global average being reported for the European Alps. As a result, the probability of climate extremes has increased and is expected to rise further. In an earlier study, we looked into immediate impacts of experimentally imposed heat waves in alpine grassland, and found that these systems were able to cope with heat as long as enough water was available. However, concomitant drought led to increased stress, and reduced aboveground biomass production and green plant cover. Here, we studied the legacy effects (lag-effects) of the imposed climate extreme to see whether delayed responses occurred and how fast the alpine grassland could rebound from the initial changes. Green cover continued to be suppressed the two following years in communities that had been exposed to the most intense hot drought, while aboveground biomass production had returned to control levels by year 2. The initial lower resistance of the forb fraction in the communities was not compensated by faster recovery later on. This resulted in alpine communities that became (and remained) relatively enriched with graminoids, which resisted the original extreme better. The responses of alpine grassland to heat extremes with or without drought observed in this study resemble those typically found in lowland grassland in the short term. However, alpine grassland exhibited longer legacy effects from an annual perspective, with delayed recovery of aboveground production and persistent changes in community composition. This suggests that once initial resistance thresholds are exceeded, impacts may be longer-lasting in alpine grassland, where recovery is constrained by both the short growing season and difficult seedling establishment.

Low temperature limits for root growth in alpine species are set by cell differentiation.

Plant growth in cold climates is not limited by carbon assimilation (source activity) but rather by reduced carbon investment into new tissues (sink limitation). It has been hypothesized that all cold-adapted plants face similar growth constraints at low temperature mainly associated with the formation of new tissues. To explore the thermal limitation of plant tissue formation, we studied root growth and anatomical root tissue characteristics in four cold-adapted alpine species (Ranunculus glacialis, Rumex alpinus, Tussilago farfara, Poa alpina), grown in thermostated soils with a vertical temperature gradient approaching 1 °C. Above-ground plant organs were exposed to typical alpine climate conditions (high solar radiation and cool nights) at 2440 m a.s.l. in the Swiss Alps to assure continuous source activity. Image-based measurements of root growth (root elongation rates at 12-h intervals, RERs) were combined with anatomical examinations in thermally constrained root tips as well as with a functional growth analysis of entire plants. Temperatures in the range 0.8 to 1.4 °C were denoted as critically low temperature thresholds for root formation across the four species. The RERs per 12 h revealed that roots kept extending at low rates at 0.7-1.2 °C but cell elongation and xylem lignification were clearly inhibited in the terminal zones of root tips. Roots exposed to temperatures between 1 and 5 °C showed strongly reduced elongation rates so that these roots contributed very little to the entire root system compared to control roots grown at 10 °C. Hardly any secondary roots were formed at temperatures below 5 °C and total root mass was substantially lower (74 % reduction in comparison to control), also the above-ground biomass was reduced by 23 %. Cell elongation and differentiation rather than cell division control length and shape of root cells at the low temperature limit of growth. Lignification of root xylem is clearly constrained at temperatures below 3 °C.

Simulated heat waves affected alpine grassland only in combination with drought.

The Alpine region is warming fast, and concurrently, the frequency and intensity of climate extremes are increasing. It is currently unclear whether alpine ecosystems are sensitive or resistant to such extremes. We subjected Swiss alpine grassland communities to heat waves with varying intensity by transplanting monoliths to four different elevations (2440-660 m above sea level) for 17 d. Half of these were regularly irrigated while the other half were deprived of irrigation to additionally induce a drought at each site. Heat waves had no significant impacts on fluorescence (Fv /Fm , a stress indicator), senescence and aboveground productivity if irrigation was provided. However, when heat waves coincided with drought, the plants showed clear signs of stress, resulting in vegetation browning and reduced phytomass production. This likely resulted from direct drought effects, but also, as measurements of stomatal conductance and canopy temperatures suggest, from increased high-temperature stress as water scarcity decreased heat mitigation through transpiration. The immediate responses to heat waves (with or without droughts) recorded in these alpine grasslands were similar to those observed in the more extensively studied grasslands from temperate climates. Responses following climate extremes may differ in alpine environments, however, because the short growing season likely constrains recovery.

Ecological consequences of the expansion of N2-fixing plants in cold biomes.

Research in warm-climate biomes has shown that invasion by symbiotic dinitrogen (N2)-fixing plants can transform ecosystems in ways analogous to the transformations observed as a consequence of anthropogenic, atmospheric nitrogen (N) deposition: declines in biodiversity, soil acidification, and alterations to carbon and nutrient cycling, including increased N losses through nitrate leaching and emissions of the powerful greenhouse gas nitrous oxide (N2O). Here, we used literature review and case study approaches to assess the evidence for similar transformations in cold-climate ecosystems of the boreal, subarctic and upper montane-temperate life zones. Our assessment focuses on the plant genera Lupinus and Alnus, which have become invasive largely as a consequence of deliberate introductions and/or reduced land management. These cold biomes are commonly located in remote areas with low anthropogenic N inputs, and the environmental impacts of N2-fixer invasion appear to be as severe as those from anthropogenic N deposition in highly N polluted areas. Hence, inputs of N from N2 fixation can affect ecosystems as dramatically or even more strongly than N inputs from atmospheric deposition, and biomes in cold climates represent no exception with regard to the risk of being invaded by N2-fixing species. In particular, the cold biomes studied here show both a strong potential to be transformed by N2-fixing plants and a rapid subsequent saturation in the ecosystem's capacity to retain N. Therefore, analogous to increases in N deposition, N2-fixing plant invasions must be deemed significant threats to biodiversity and to environmental quality.

Plant species dominance shifts across erosion edge-meadow transects in the Swiss Alps.

While exerting no obvious function under "average" environmental conditions, the presence of certain plant specialists becomes crucial in the event of a complete failure of a community due to severe disturbance such as landslides. Plants capable of growing at erosion edges may act as potential edge-engineers by coping with unstable ground and stabilizing the soil with their roots. We hypothesized that life conditions at erosion edges select for a particular set of specialists or species with specific traits, the identification of which was the aim of the study. Across 17 small-scale transects (0.40 × 1.60 m) from intact meadows to landslide edges (Ursern Valley, Swiss Alps, c. 1,600 m a.s.l.), we quantified plant species abundance by the point intercept method and characterized growth conditions based on Landolt's indicator values, leaf δ(13)C, and volumetric soil moisture in the uppermost soil layers. We observed a clear change of plant species composition and relative abundance from the meadow to the edge, presumably induced by the 25 % lower soil moisture and microclimatic exposure. Species richness at the edge was two-thirds of that in the meadow, but was positively correlated with species richness of the adjacent meadow. Species with "edge-preference" had either (1) rolled or festucoid leaves like Festuca spp., Avenella flexuosa and Nardus stricta, or (2) small, scleromorphic leaves like Vaccinium vitis-idaea, Calluna vulgaris and Thymus ssp. Graminoids with rolled/festucoid leaves were found to be the most dominant edge-specialists. The grass Festuca valesiaca s.l. emerged as the most dominant plant species at the edge, having an 11-times higher cover at the edge than in the meadow. In this montane grassland, a single species contributes to the stabilization of erosion edges and may be regarded as a potential keystone species for slope stability and regeneration after landslides even its role has not so far been established.

Manganese deficiency of silver fir trees (Abies alba) at a reforested site in the Jura mountains, Switzerland: aspects of cause and effect.

We examined manganese uptake and translocation in 30-year-old silver fir trees (Abies alba Mill.) showing severe symptoms of needle chlorosis by analyzing both xylem and phloem sap of shoots and an extract of root sap originating from both xylem and phloem elements. Manganese concentrations in shoot xylem and phloem sap were significantly lower in chlorotic trees than in neighboring healthy trees. The Mn concentration of root sap was also lower in these Mn-deficient trees than in healthy trees, indicating reduced Mn uptake by Mn-deficient trees. Although Mn deficiency had no effect on the Mn concentration of whole roots, separation of root tissue into root cortex and stele (wood) suggested impaired translocation of Mn from the cortex to the stele in Mn-deficient trees. Triphenyltetrazolium chloride (TTC) tests indicated that there was no decrease in reducing capacity of the fine roots (< 1 mm in diameter) of Mn-deficient trees. Fine roots of Mn-deficient trees contained significantly more organic acids than fine roots of healthy trees, including increased concentrations of malic, quinic, trans-acontic and formic acid; however, concentrations of pyruvic and acetic acid were lower than in fine roots of healthy trees. The total amounts of organic acids in the rhizospheric soil were similar for healthy and Mn-deficient trees. Pyruvic acid concentration was significantly increased in the rhizospheric soil of Mn-deficient trees, and concentrations of simple aliphatic acids like formic and acetic acid also tended to be higher in the rhizospheric soil of Mn-deficient trees. Both pH and water content were higher in bulk soil and rhizospheric soil around Mn-deficient trees compared with soil around healthy trees. Although lower concentrations of exchangeable Mn were found in the soil around Mn-deficient trees, the active Mn concentration (sum of exchangeable and reducible Mn) did not differ between healthy and Mn-deficient trees. A considerable proportion of manganese was in an oxidized form in the soil around Mn-deficient trees.