Dampening effects of long‐term experimental drought on growth and mortality rates of a Holm oak forest

Forests respond to increasing intensities and frequencies of drought by reducing growth and with higher tree mortality rates. Little is known, however, about the long‐term consequences of generally drier conditions and more frequent extreme droughts. A Holm oak forest was exposed to experimental rainfall manipulation for 13 years to study the effect of increasing drought on growth and mortality of the dominant species Quercus ilex, Phillyrea latifolia, and Arbutus unedo. The drought treatment reduced stem growth of A. unedo (?66.5%) and Q. ilex (?17.5%), whereas P. latifolia remained unaffected. Higher stem mortality rates were noticeable in Q. ilex (+42.3%), but not in the other two species. Stem growth was a function of the drought index of early spring in the three species. Stem mortality rates depended on the drought index of winter and spring for Q. ilex and in spring and summer for P. latifolia, but showed no relation to climate in A. unedo. Following a long and intense drought (2005–2006), stem growth of Q. ilex and P. latifolia increased, whereas it decreased in A. unedo. Q. ilex also enhanced its survival after this period. Furthermore, the effect of drought treatment on stem growth in Q. ilex and A. unedo was attenuated as the study progressed. These results highlight the different vulnerabilities of Mediterranean species to more frequent and intense droughts, which may lead to partial species substitution and changes in forest structure and thus in carbon uptake. The response to drought, however, changed over time. Decreased intra‐ and interspecific competition after extreme events with high mortality, together with probable morphological and physiological acclimation to drought during the study period, may, at least in the short term, buffer forests against drier conditions. The long‐term effects of drought consequently deserve more attention, because the ecosystemic responses are unlikely to be stable over time.Nontechnical summaryIn this study, we evaluate the effect of long‐term (13 years) experimental drought on growth and mortality rates of three forest Mediterranean species, and their response to the different intensities and durations of natural drought. We provide evidence for species‐specific responses to drought, what may eventually lead to a partial community shift favoring the more drought‐resistant species. However, we also report a dampening of the treatment effect on the two drought‐sensitive species, which may indicate a potential adaptation to drier conditions at the ecosystem or population level. These results are thus relevant to account for the stabilizing processes that would alter the initial response of ecosystem to drought through changes in plant physiology, morphology, and demography compensation.     

Enhanced evapotranspiration was observed during extreme drought from Miscanthus,opposite of other crops

The impact of extreme drought and heat stress that occurred in the Midwestern U.S. in 2012 on evapotranspiration (ET), net ecosystem productivity (NEP), and water‐use efficiency (WUE) of three perennial ecosystems (switchgrass, miscanthus, prairie) and a maize/soybean agroecosystem was studied as part of a long‐term experiment. Miscanthus had a slower initial response but an eventually drastic ET as drought intensified, which resulted in the largest water deficit among the crops. The substantially higher ET at peak drought was likely supplied by access to deep soil water, but suggests that stomatal conductance of miscanthus during the drought may respond differently than the other ecosystems, consistent with an anisohydric strategy. While there was a discrepancy in the water consumption of maize and switchgrass/prairie in the early time of drought, all these ecosystems followed a water‐saving strategy when drought intensified. The gross primary production (GPP) of miscanthus dropped, but was reversible, when temperature reached 40 °C and still provided the largest total GPP among the ecosystems. Increased ET for miscanthus during 2012 resulted a large decline in ecosystem WUE compared to what was observed in other years. The biophysical responses of miscanthus measured during an extreme, historic drought suggest that this species can maintain high productivity longer than other ecosystems during a drought at the expense of water use. While miscanthus maintained productivity during drought, recovery lagged associated with depleted soil moisture. The enhanced ET of miscanthus may intensify droughts through increase supply of deep soil moisture to the atmosphere.     

Plant‐mediated effects of drought on aphid population structure and parasitoid attack

The effects of predicted climate change on aphid–natural enemy interactions have principally considered the effects of elevated carbon dioxide concentration and air temperature. However, increased incidence of summer droughts are also predicted in Northern Europe, which could affect aphid–plant interactions and aphid antagonists. We investigated how simulated summer drought affected the bird cherry–oat aphid, Rhopalosiphum padi L., and its natural enemy the parasitoid wasp Aphidius ervi. Drought and, to a greater extent, aphids reduced barley ( Hordeum vulgare) dry mass by 33% and 39%, respectively. Drought reduced leaf and root nitrogen concentrations by 13% and 28%, respectively, but foliar amino acid concentrations and composition remained similar. Aphid numbers were unaffected by drought, but population demography changed significantly; adults constituted 41% of the population on drought‐treated plants, but only 26% on those receiving ambient irrigation. Nymphs constituted 56% and 69% of the population on these plants, respectively, suggesting altered aphid development rates on drought‐stressed plants. Parasitism rates were significantly lower on drought‐stressed plants (9 attacks h^?1 compared with 35 attacks h^?1 on ambient‐irrigated plants), most likely because of lower incidence of nymphs and more adults, the latter being more difficult to parasitize. Any physiological changes in individual aphids did not affect parasitoid preferences, suggesting that attacks were postponed because of drought‐induced changes in aphid demography. This study demonstrates the potential for sporadic climate change events, such as summer drought, to be disruptive to herbivore–antagonist interactions.     

The temporal response to drought in a Mediterranean evergreen tree: comparing a regional precipitation gradient and a throughfall exclusion experiment

Like many midlatitude ecosystems, Mediterranean forests will suffer longer and more intense droughts with the ongoing climate change. The responses to drought in long‐lived trees differ depending on the time scale considered, and short‐term responses are currently better understood than longer term acclimation. We assessed the temporal changes in trees facing a chronic reduction in water availability by comparing leaf‐scale physiological traits, branch‐scale hydraulic traits, and stand‐scale biomass partitioning in the evergreen Quercus ilex across a regional precipitation gradient (long‐term changes) and in a partial throughfall exclusion experiment (TEE, medium term changes). At the leaf scale, gas exchange, mass per unit area and nitrogen concentration showed homeostatic responses to drought as they did not change among the sites of the precipitation gradient or in the experimental treatments of the TEE. A similar homeostatic response was observed for the xylem vulnerability to cavitation at the branch scale. In contrast, the ratio of leaf area over sapwood area (LA/SA) in young branches exhibited a transient response to drought because it decreased in response to the TEE the first 4 years of treatment, but did not change among the sites of the gradient. At the stand scale, leaf area index (LAI) decreased, and the ratios of stem SA to LAI and of fine root area to LAI both increased in trees subjected to throughfall exclusion and from the wettest to the driest site of the gradient. Taken together, these results suggest that acclimation to chronic drought in long‐lived Q. ilex is mediated by changes in hydraulic allometry that shift progressively from low (branch) to high (stand) organizational levels, and act to maintain the leaf water potential within the range of xylem hydraulic function and leaf photosynthetic assimilation.     

Growth and resilience responses of Scots pine to extreme droughts across Europe depend on predrought growth conditions

Global climate change is expected to further raise the frequency and severity of extreme events, such as droughts. The effects of extreme droughts on trees are difficult to disentangle given the inherent complexity of drought events (frequency, severity, duration, and timing during the growing season). Besides, drought effects might be modulated by trees’ phenotypic variability, which is, in turn, affected by long‐term local selective pressures and management legacies. Here we investigated the magnitude and the temporal changes of tree‐level resilience (i.e., resistance, recovery, and resilience) to extreme droughts. Moreover, we assessed the tree‐, site‐, and drought‐related factors and their interactions driving the tree‐level resilience to extreme droughts. We used a tree‐ring network of the widely distributed Scots pine (Pinus sylvestris) along a 2,800 km latitudinal gradient from southern Spain to northern Germany. We found that the resilience to extreme drought decreased in mid‐elevation and low productivity sites from 1980–1999 to 2000–2011 likely due to more frequent and severe droughts in the later period. Our study showed that the impact of drought on tree‐level resilience was not dependent on its latitudinal location, but rather on the type of sites trees were growing at and on their growth performances (i.e., magnitude and variability of growth) during the predrought period. We found significant interactive effects between drought duration and tree growth prior to drought, suggesting that Scots pine trees with higher magnitude and variability of growth in the long term are more vulnerable to long and severe droughts. Moreover, our results indicate that Scots pine trees that experienced more frequent droughts over the long‐term were less resistant to extreme droughts. We, therefore, conclude that the physiological resilience to extreme droughts might be constrained by their growth prior to drought, and that more frequent and longer drought periods may overstrain their potential for acclimation.     

Species‐specific differences in water uptake depth of mature temperate trees vary with water availability in the soil

• Temperate tree species differ in their physiological sensitivity to declining soil moisture and drought. Although species‐specific responses to drought have often been suggested to be the result of different water uptake depths, empirical evidence for such a mechanism is scarce.
• Here we test if differences in water uptake depths can explain previously observed species‐specific physiological responses of temperate trees to drought and if the water uptake depth of different species varies in response to declining soil moisture. For this purpose, we employed stable oxygen and hydrogen isotopes of soil and xylem water that we collected over the course of three growing seasons in a mature temperate forest in Switzerland.
• Our data show that all investigated species utilise water from shallow soil layers during times of sufficient soil water supply. However, Fraxinus excelsior, Fagus sylvatica and Acer pseudoplatanus were able to shift their water uptake to deeper soil layers when soil water availability decreased in the topsoil. In contrast, Picea abies, was not able to shift its water uptake to deeper soil layers.
• We conclude from our data that more drought‐resistant tree species are able to shift their water uptake to deeper soil layers when water availability in the topsoil is becoming scarce. In addition, we were able to show that water uptake depth of temperate tree species is a trait with high plasticity that needs to be characterised across a range of environmental conditions.

     

Quantitative losses vs. qualitative stability of ectomycorrhizal community responses to 3 years of experimental summer drought in a beech–spruce forest

Forest ecosystems in central Europe are predicted to face an increasing frequency and severity of summer droughts because of global climate change. European beech and Norway spruce often coexist in these forests with mostly positive effects on their growth. However, their different below‐ground responses to drought may lead to differences in ectomycorrhizal (ECM) fungal community composition and functions which we examined at the individual root and ecosystem levels. We installed retractable roofs over plots in Kranzberg Forest (11°39′42″E, 48°25′12″N; 490 m a.s.l.) to impose repeated summer drought conditions and assigned zones within each plot where trees neighboured the same or different species to study mixed species effects. We found that ECM fungal community composition changed and the numbers of vital mycorrhizae decreased for both tree species over 3 drought years (2014–2016), with the ECM fungal community diversity of beech exhibiting a faster and of spruce a stronger decline. Mixed stands had a positive effect on the ECM fungal community diversity of both tree species after the third drought year. Ectomycorrhizae with long rhizomorphs increased in both species under drought, indicating long‐distance water transport. However, there was a progressive decline in the number of vital fine roots during the experiment, resulting in a strong reduction in enzyme activity per unit volume of soil. Hydrolytic enzyme activities of the surviving ectomycorrhizae were stable or stimulated upon drought, but there was a large decline in ECM fungal species with laccase activity, indicating a decreased potential to exploit nutrients bound to phenolic compounds. Thus, the ectomycorrhizae responded to repeated drought by maintaining or increasing their functionality at the individual root level, but were unable to compensate for quantitative losses at the ecosystem level. These findings demonstrate a strong below‐ground impact of recurrent drought events in forests.     

Overexpression of ubiquitin‐like LpHUB1 gene confers drought tolerance in perennial ryegrass

HUB1, also known as Ubl5, is a member of the subfamily of ubiquitin‐like post‐translational modifiers. HUB1 exerts its role by conjugating with protein targets. The function of this protein has not been studied in plants. A HUB1 gene, LpHUB1, was identified from serial analysis of gene expression data and cloned from perennial ryegrass. The expression of this gene was reported previously to be elevated in pastures during the summer and by drought stress in climate‐controlled growth chambers. Here, pasture‐type and turf‐type transgenic perennial ryegrass plants overexpressing LpHUB1 showed improved drought tolerance, as evidenced by improved turf quality, maintenance of turgor and increased growth. Additional analyses revealed that the transgenic plants generally displayed higher relative water content, leaf water potential, and chlorophyll content and increased photosynthetic rate when subjected to drought stress. These results suggest HUB1 may play an important role in the tolerance of perennial ryegrass to abiotic stresses.     

Down‐regulation of CBP80 gene expression as a strategy to engineer a drought‐tolerant potato

Developing new strategies for crop plants to respond to drought is crucial for their innovative breeding. The down‐regulation of nuclear cap‐binding proteins in Arabidopsis renders plants drought tolerant. The CBP80 gene in the potato cultivar Desiree was silenced using artificial microRNAs. Transgenic plants displayed a higher tolerance to drought, ABA‐hypersensitive stomatal closing, an increase in leaf stomata and trichome density, and compact cuticle structures with a lower number of microchannels. These findings were correlated with a higher tolerance to water stress. The level of miR159 was decreased, and the levels of its target mRNAs MYB33 and MYB101 increased in the transgenic plants subjected to drought. Similar trends were observed in an Arabidopsis cbp80 mutant. The evolutionary conservation of CBP80, a gene that plays a role in the response to drought, suggests that it is a candidate for genetic manipulations that aim to obtain improved water‐deficit tolerance of crop plants.     

Effects of drought on fish across axes of space, time and ecological complexity

• 1 We evaluate the position of 50 previously published studies of fish and drought with respect to spatial scale of study (individual stream pools to subcontinents), length of the dry period (weeks to centuries), and level of system complexity (individual fish to ecosystems). Most papers address short (months to a year) droughts or dry periods, in local reaches of streams, and impacts on populations or local assemblages. In these 50 papers, the most frequently demonstrated effects of drought were population declines, loss of habitat, changes in the community, negative effects from changes in water quality, movement within catchments, and crowding of fish in reduced microhabitats. Thirteen other less frequent effects also were identified.
• 2 Gaps in knowledge exist on effects of long‐term droughts (decades to centuries), influence of drought on fish effects in ecosystems, and at the spatial scale of river basins to subcontinents. However, some of these gaps have recently been addressed, particularly additive effects of repeated drying episodes and whole‐lake or basin‐wide effects of drought, and in using molecular techniques to seek signals of drought at wide geographic scales because of events in the deep past. Gaps in knowledge remain for effects of very short dry periods, on drought effects on higher levels of complexity, and on the manner in which droughts at the scale of decades affect fish.
• 3 Data from streams in Oklahoma and elsewhere in the south‐western U.S.A. suggest that most droughts may leave little persistent signal in the existing fish fauna, i.e. that recovery from drought by fish populations or assemblages in the region can be rapid. However, species that are vulnerable to drought or water loss in streams may have disappeared from some basins in the region before the mid‐1900s, and recent evidence also suggests that extreme droughts do sometimes alter fish assemblages.
• 4 Little is known about mechanisms by which droughts have direct or indirect effects on fish, the roles of droughts in the evolution of fish species, and the ways droughts alter effects of fish in ecosystems. Global climate changes may have serious consequences for future local or regional fish faunas, but ongoing studies of fish experiencing drought may aid in future conservation of what will become species at risk under climate‐change scenarios.

     

Shift in community structure in an early‐successional Mediterranean shrubland driven by long‐term experimental warming and drought and natural extreme droughts

Global warming and recurring drought are expected to accelerate water limitation for plant communities in semiarid Mediterranean ecosystems and produce directional shifts in structure and composition that are not easily detected, and supporting evidence is scarce. We conducted a long‐term (17 years) nocturnal‐warming (+0.6°C) and drought (?40% rainfall) experiments in an early‐successional Mediterranean shrubland to study the changes in community structure and composition, contrasting functional groups and dominant species, and the superimposed effects of natural extreme drought. Species richness decreased in both the warming and drought treatments. Responses to the moderate warming were associated with decreases in herb abundance, and responses to the drought were associated with decreases in both herb and shrub abundances. The drought also significantly decreased community diversity and evenness. Changes in abundance differed between herbs (decreases) and shrubs (increases or no changes). Both warming and drought, especially drought, increased the relative species richness and abundance of shrubs, favoring the establishment of shrubs. Both warming and drought produced significant shifts in plant community composition. Experimental warming shifted the community composition from Erica multiflora toward Rosmarinus officinalis, and drought consistently shifted the composition toward Globularia alypum. The responses in biodiversity (e.g., community biodiversity, changes of functional groups and compositional shifts) were also strongly correlated with atmospheric drought (SPEI) in winter–spring and/or summer, indicating sensitivity to water limitation in this early‐successional Mediterranean ecosystem, especially to natural extreme droughts. Our results suggest that the shifts in species assembles and community diversity and composition are accelerated by the long‐term nocturnal‐warming and drought, combined with natural severe droughts, and that the magnitude of the impacts of climate change is also correlated with the successional status of ecosystem. The results thus highlight the necessity for assessing the impacts on ecosystemic functioning and services and developing effective measures for conserving biodiversity.     

Drought footprint on European ecosystems between 1999 and 2010 assessed by remotely sensed vegetation phenology and productivity

Drought affects more people than any other natural disaster but there is little understanding of how ecosystems react to droughts. This study jointly analyzed spatio‐temporal changes of drought patterns with vegetation phenology and productivity changes between 1999 and 2010 in major European bioclimatic zones. The Standardized Precipitation and Evapotranspiration Index (SPEI) was used as drought indicator whereas changes in growing season length and vegetation productivity were assessed using remote sensing time‐series of Normalized Difference Vegetation Index (NDVI). Drought spatio‐temporal variability was analyzed using a Principal Component Analysis, leading to the identification of four major drought events between 1999 and 2010 in Europe. Correspondence Analysis showed that at the continental scale the productivity and phenology reacted differently to the identified drought events depending on ecosystem and land cover. Northern and Mediterranean ecosystems proved to be more resilient to droughts in terms of vegetation phenology and productivity developments. Western Atlantic regions and Eastern Europe showed strong agglomerations of decreased productivity and shorter vegetation growing season length, indicating that these ecosystems did not buffer the effects of drought well. In a climate change perspective, increase in drought frequency or intensity may result in larger impacts over these ecosystems, thus management and adaptation strategies should be strengthened in these areas of concerns.     

Shifts in population size structure for a drying‐tolerant fish in response to extreme drought

For freshwater systems, climate change‐induced alterations to drought regimes are a considerable threat to already threatened species. This is particularly poignant for kōwaro (or Canterbury mudfish, Neochanna burrowsius), a critically endangered fish largely restricted to drying‐prone waterways on the Canterbury Plains, New Zealand. By comparing three catchment‐wide surveys (2007, 2010, 2015) within the Waianiwaniwa Valley, we assessed the scale and magnitude of population change induced by 2 years of consecutive drought (2014–15), when compared to surveys during wetter conditions (2007, 2010). The droughts triggered a catchment‐wide switch from adult‐dominated populations to populations comprised of juveniles indicated by a significant reduction in median size (~95 mm during the wet to ~60 mm after drought). In comparison, population abundances were highly variable, indicated by no significant change in catch‐per‐unit‐effort. The large variation in catch rates and connection of median size to reproductive potential mean median size will be useful to measure in monitoring to infer potential changes to population resilience, particularly during extreme events. Furthermore, because N. burrowsius could be regarded as extremophile fish, already restricted to harsh habitats, they are likely to become increasingly threatened by climate change. Thus, tools that allow for insightful comparisons between populations, such as a population resilience framework based on both abundance and body size distribution, will be increasingly important for pragmatic decision‐making for targeted conservation measures.     

Tree diversity mitigates defoliation after a drought‐induced tipping point

Understanding the processes that underlie drought‐related tree vitality loss is essential for anticipating future forest dynamics, and for developing management plans aiming at increasing the resilience of forests to climate change. Forest vitality has been continuously monitored in Europe since the acid rain alert in the 1980s, and the intensive monitoring plots of ICP Forests offer the opportunity to investigate the effects of air pollution and climate change on forest condition. By making use of over 100 long‐term monitoring plots, where crown defoliation has been assessed extensively since 1990, we discovered a progressive shift from a negative to a positive effect of species richness on forest health. The observed tipping point in the balance of net interactions, from competition to facilitation, has never been reported from real ecosystems outside experimental conditions; and the strong temporal consistency of our observations with increasing drought stress emphasizes its climate change relevance. Furthermore, we show that higher species diversity has reduced the severity of defoliation in the long term. Our results confirm the greater resilience of diverse forests to future climate change‐induced stress. More generally, they add to an accumulating body of evidence on the large potential of tree species mixtures to face manifold disturbances in a changing world.     

Community structures of N2‐fixing bacteria associated with the phyllosphere of a Holm oak forest and their response to drought

Biological nitrogen (N) fixation is a key pathway in terrestrial ecosystems and is therefore critical for understanding the responses of ecosystems to global environmental changes. The free‐living diazotrophic community is distributed along the canopy‐to‐soil profile, but the ecological significance of epiphyllic N₂ fixers, despite their functional relevance, on plant foliar surfaces remains very poorly understood compared with the N₂‐fixing community in forest litter and soils. We assessed the community structure of N₂ fixers and overall bacteria by genetic fingerprinting (t‐RFLP) to explore the seasonal successional patterns of the microbial community in the natural phyllosphere of a Holm oak (Quercus ilex) forest submitted to 12‐year field experiment of rain exclusion mimicking the conditions of drought projected for the coming decades. Leaves of Holm oak were analysed in different seasons over a period of 1.5 years. The bacterial community of the phyllosphere did not correspond to the surrounding soil biome in the same area. These analyses provided field evidence for the presence of free‐living diazotrophs associated with the tissues of leaves of Holm oak, the dominant tree species of many Mediterranean forests. The results also revealed that the community composition is affected seasonally and inter‐annually by the environment, and that the composition shifts in response to climate change. Drought treatment increased the richness of the epiphyllic microbial community, especially during the summer. These changes were associated with higher C:N ratios of leaves observed in response to drought in semiarid areas. This epiphyllic microbiota that can potentially fix N₂ extends the capacity of plants to adapt to the environment.     

Expression of the CCCH‐tandem zinc finger protein gene OsTZF5 under a stress‐inducible promoter mitigates the effect of drought stress on rice grain yield under field conditions

Increasing drought resistance without sacrificing grain yield remains an ongoing challenge in crop improvement. In this study, we report that O ryza s ativa CCCH‐t andem z inc f inger protein 5 (OsTZF5) can confer drought resistance and increase grain yield in transgenic rice plants. Expression of OsTZF5 was induced by abscisic acid, dehydration and cold stress. Upon stress, OsTZF5‐GFP localized to the cytoplasm and cytoplasmic foci. Transgenic rice plants overexpressing OsTZF5 under the constitutive maize ubiquitin promoter exhibited improved survival under drought but also growth retardation. By introducing OsTZF5 behind the stress‐responsive OsNAC6 promoter in two commercial upland cultivars, Curinga and NERICA4, we obtained transgenic plants that showed no growth retardation. Moreover, these plants exhibited significantly increased grain yield compared to non‐transgenic cultivars in different confined field drought environments. Physiological analysis indicated that OsTZF5 promoted both drought tolerance and drought avoidance. Collectively, our results provide strong evidence that OsTZF5 is a useful biotechnological tool to minimize yield losses in rice grown under drought conditions.     

Impacts of climatic stability on the structural and functional aspects of macroinvertebrate communities after severe drought

1. Unpredictable, extreme climatic events (e.g. droughts) can potentially destabilize aquatic communities. From 1998 to 2002, southwestern Georgia, U.S.A., experienced the third worst drought of the last 100 years, leading to loss of surface flow in many small streams. We sampled macroinvertebrates, flow and water chemistry in small headwater streams from 2001 to 2007 in two adjacent coastal plain streams of contrasting headwater type (wetland and groundwater‐fed seep) following resumption of flow. 2. Decreasing water temperature, conductivity and nutrient concentrations within the first 2 years of the study indicated flushing of the streambed associated with increased discharge. Invertebrate community composition became less variable over time and during wetter years, reflecting water chemistry, hydrological and climate conditions. 3. A core set of species appeared immediately following breaking of the drought in both streams, reflecting a shared species pool. These species exhibited resilience traits, including short life cycles and resistance to desiccation, which allowed for rapid recovery from disturbance. Such species, which were small‐bodied, sclerotized and abundant in the drift, were then replaced as flows increased by those that were larger, soft‐bodied and rare in drift, suggesting a more stable and less ephemeral habitat. 4. Hydrologic regime and long‐term precipitation indices were strongly correlated with invertebrate community and trait structure. Long‐term data allowed for better interpretation of the effects of infrequent disturbances on aquatic ecosystems. Additionally, long‐term precipitation indices (i.e. 48‐month standardized precipitation index) can indicate the likelihood of a return to drought, allowing for the collection of pre‐disturbance data.     

Not all droughts are created equal: the impacts of interannual drought pattern and magnitude on grassland carbon cycling

Climate extremes, such as drought, may have immediate and potentially prolonged effects on carbon cycling. Grasslands store approximately one‐third of all terrestrial carbon and may become carbon sources during droughts. However, the magnitude and duration of drought‐induced disruptions to the carbon cycle, as well as the mechanisms responsible, remain poorly understood. Over the next century, global climate models predict an increase in two types of drought: chronic but subtle ‘press‐droughts’, and shorter term but extreme ‘pulse‐droughts’. Much of our current understanding of the ecological impacts of drought comes from experimental rainfall manipulations. These studies have been highly valuable, but are often short term and rarely quantify carbon feedbacks. To address this knowledge gap, we used the Community Land Model 4.0 to examine the individual and interactive effects of pulse‐ and press‐droughts on carbon cycling in a mesic grassland of the US Great Plains. A series of modeling experiments were imposed by varying drought magnitude (precipitation amount) and interannual pattern (press‐ vs. pulse‐droughts) to examine the effects on carbon storage and cycling at annual to century timescales. We present three main findings. First, a single‐year pulse‐drought had immediate and prolonged effects on carbon storage due to differential sensitivities of ecosystem respiration and gross primary production. Second, short‐term pulse‐droughts caused greater carbon loss than chronic press‐droughts when total precipitation reductions over a 20‐year period were equivalent. Third, combining pulse‐ and press‐droughts had intermediate effects on carbon loss compared to the independent drought types, except at high drought levels. Overall, these results suggest that interannual drought pattern may be as important for carbon dynamics as drought magnitude and that extreme droughts may have long‐lasting carbon feedbacks in grassland ecosystems.     

Impact of defoliation intensities on plant biomass,nutrient uptake and arbuscular mycorrhizal symbiosis in Lotus tenuis growing in a saline‐sodic soil

The impact of different defoliation intensities on the ability of Lotus tenuis plants to regrowth, mobilise nutrients and to associate with native AM fungi and Rhizobium in a saline‐sodic soil was investigated. After 70 days, plants were subjected to 0, 25, 50, 75 and 100% defoliation and shoot regrowth was assessed at the end of subsequent 35 days. Compared to non‐defoliated plants, low or moderate defoliation up to 75% did not affect shoot regrowth. However, 100% treatment affected shoot regrowth and the clipped plants were not able to compensate the growth attained by non‐defoliated plants. Root growth was more affected by defoliation than shoot growth. P and N concentrations in shoots and roots increased with increasing defoliation while Na⁺ concentration in shoots of non‐defoliated and moderately defoliated plants was similar. Non‐defoliated and moderately defoliated plants prevented increases of Na⁺ concentration in shoots through both reducing Na⁺ uptake and Na⁺ transport to shoots by accumulating Na⁺ in roots. At high defoliation, the salinity tolerance mechanism is altered and Na⁺ concentration in shoots was higher than in roots. Reduction in the photosynthetic capacity induced by defoliation neither changed the root length colonised by AM fungi nor arbuscular colonisation but decreased the vesicular colonisation. Spore density did not change, but hyphal density and Rhizobium nodules increased with defoliation. The strategy of the AM symbiont consists in investing most of the C resources to preferentially retain arbuscular colonisation as well as inoculum density in the soil.