Enhanced growth resistance but no decline in growth resilience under long-term extreme droughts.

The frequency, intensity, and duration of extreme droughts, with devastating impacts on tree growth and survival, have increased with climate change over the past decades. Assessing growth resistance and resilience to drought is a crucial prerequisite for understanding the responses of forest functioning to drought events. However, the responses of growth resistance and resilience to extreme droughts with different durations across different climatic zones remain unclear. Here, we investigated the spatiotemporal patterns in growth resistance and resilience in response to extreme droughts with different durations during 1901-2015, relying on tree-ring chronologies from 2389 forest stands over the mid- and high-latitudinal Northern Hemisphere, species-specific plant functional traits, and diverse climatic factors. The findings revealed that growth resistance and resilience under 1-year droughts were higher in humid regions than in arid regions. Significant higher growth resistance was observed under 2-year droughts than under 1-year droughts in both arid and humid regions, while growth resilience did not show a significant difference. Temporally, tree growth became less resistant and resilient to 1-year droughts in 1980-2015 than in 1901-1979 in both arid and humid regions. As drought duration lengthened, the predominant impacts of climatic factors on growth resistance and resilience weakened and instead foliar economic traits, plant hydraulic traits, and soil properties became much more important in both climatic regions; in addition, such trends were also observed temporally. Finally, we found that most of the Earth system models (ESMs) used in this study overestimated growth resistance and underestimated growth resilience under both 1-year and 2-year droughts. A comprehensive ecophysiological understanding of tree growth responses to longer and intensified drought events is urgently needed, and a specific emphasis should be placed on improving the performance of ESMs.

Nature-based framework for sustainable afforestation in global drylands under changing climate.

Drylands cover more than 40% of Earth's land surface and occur at the margin of forest distributions due to the limited availability of water for tree growth. Recent elevated temperature and low precipitation have driven greater forest declines and pulses of tree mortality on dryland sites compared to humid sites, particularly in temperate Eurasia and North America. Afforestation of dryland areas has been widely implemented and is expected to increase in many drylands globally to enhance carbon sequestration and benefits to the human environment, but the interplay of sometimes conflicting afforestation outcomes has not been formally evaluated yet. Most previous studies point to conflicts between additional forest area and water consumption, in particular water yield and soil conservation/desalinization in drylands, but were generally confined to local and regional scales. Our global synthesis demonstrates that additional tree cover can amplify water consumption through a nonlinear increase in evapotranspiration-depending on tree species, age, and structure-which will be further intensified by future climate change. In this review we identify substantial knowledge gaps in addressing the dryland afforestation dilemma, where there are trade-offs with planted forests between increased availability of some resources and benefits to human habitats versus the depletion of other resources that are required for sustainable development of drylands. Here we propose a method of addressing comprehensive vegetation carrying capacity, based on regulating the distribution and structure of forest plantations to better deal with these trade-offs in forest multifunctionality. We also recommend new priority research topics for dryland afforestation, including: responses and feedbacks of dryland forests to climate change; shifts in the ratio of ecosystem ET to tree cover; assessing the role of scale of afforestation in influencing the trade-offs of dryland afforestation; and comprehensive modeling of the multifunctionality of dryland forests, including both ecophysiological and socioeconomic aspects, under a changing climate.

Seasonal variations in water flux compositions controlled by leaf development: isotopic insights at the canopy-atmosphere interface.

Water-stable isotopes provide a valuable tool for tracing plant-water interactions, particularly evapotranspiration (ET) partitioning and leaf water dynamics at the plant-atmosphere interface. However, process-based investigations of plant/leaf development and the associated isotopic dynamics of water fluxes involving isotope enrichment at plant-atmosphere interfaces at the ecosystem scale remain challenging. In this study, in situ isotopic measurements and tracer-aided models were used to study the dynamic interactions between vegetation growth and the isotopic dynamics of water fluxes (ET, soil evaporation, and transpiration) involving isotope enrichment in canopy leaves in a multispecies grassland ecosystem. The day-to-day variations in the isotopic compositions of ET flux were mainly controlled by plant growth, which could be explained by the significant logarithmic relationship determined between the leaf area index and transpiration fraction. Leaf development promoted a significant increase in the isotopic composition of ET and led to a slight decrease in the isotopic composition of water in canopy leaves. The transpiration (evaporation) isoflux acted to increase (decrease) the δ18O of water vapor, and the total isoflux impacts depended on the seasonal tradeoffs between transpiration and evaporation. The isotopic evidence in ET fluxes demonstrates the biotic controls on day-to-day variations in water/energy flux partitioning through transpiration activity. This study emphasizes that stable isotopes of hydrogen and oxygen are effective tools for quantitative evaluations of the hydrological component partitioning of ecosystems and plant-climate interactions.

Uneven winter snow influence on tree growth across temperate China.

Winter snow is an important driver of tree growth in regions where growing-season precipitation is limited. However, observational evidence of this influence at larger spatial scales and across diverse bioclimatic regions is lacking. Here, we investigated the interannual effects of winter (here defined as previous October to current February) snow depth on tree growth across temperate China over the period of 1961-2015, using a regional network of tree ring records, in situ daily snow depth observations, and gridded climate data. We report uneven effects of winter snow depth on subsequent growing-season tree growth across temperate China. There shows little effect on tree growth in drier regions that we attribute mainly to limited snow accumulation during winter. By contrast, winter snow exerts important positive influence on tree growth in stands with high winter snow accumulation (e.g., in parts of cold arid regions). The magnitude of this effect depends on the proportion of winter snow to pre-growing-season (previous October to current April) precipitation. We further observed that tree growth in drier regions tends to be increasingly limited by warmer growing-season temperature and early growing-season water availability. No compensatory effect of winter snow on the intensifying drought limitation of tree growth was observed across temperate China. Our findings point toward an increase in drought vulnerability of temperate forests in a warming climate.

Divergent evapotranspiration partition dynamics between shrubs and grasses in a shrub-encroached steppe ecosystem.

Previous evapotranspiration (ET) partitioning studies have usually neglected competitions and interactions between antagonistic plant functional types. This study investigated whether shrubs and grasses have divergent ET partition dynamics impacted by different water-use patterns, canopy structures, and physiological properties in a shrub-encroached steppe ecosystem in Inner Mongolia, China. The soil water-use patterns of shrubs and grasses have been quantified by an isotopic tracing approach and coupled into an improved multisource energy balance model to partition ET fluxes into soil evaporation, grass transpiration, and shrub transpiration. The mean fractional contributions to total ET were 24 ± 13%, 20 ± 4%, and 56 ± 16% for shrub transpiration, grass transpiration, and soil evaporation respectively during the growing season. Difference in ecohydrological connectivity and leaf development both contributed to divergent transpiration partitioning between shrubs and grasses. Shrub-encroachment processes result in larger changes in the ET components than in total ET flux, which could be well explained by changes in canopy resistance, an ecosystem function dominated by the interaction of soil water-use patterns and ecosystem structure. The analyses presented here highlight the crucial effects of vegetation structural changes on the processes of land-atmosphere interaction and climate feedback.

Differentiating drought legacy effects on vegetation growth over the temperate Northern Hemisphere.

In view of future changes in climate, it is important to better understand how different plant functional groups (PFGs) respond to warmer and drier conditions, particularly in temperate regions where an increase in both the frequency and severity of drought is expected. The patterns and mechanisms of immediate and delayed impacts of extreme drought on vegetation growth remain poorly quantified. Using satellite measurements of vegetation greenness, in-situ tree-ring records, eddy-covariance CO2 and water flux measurements, and meta-analyses of source water of plant use among PFGs, we show that drought legacy effects on vegetation growth differ markedly between forests, shrubs and grass across diverse bioclimatic conditions over the temperate Northern Hemisphere. Deep-rooted forests exhibit a drought legacy response with reduced growth during up to 4 years after an extreme drought, whereas shrubs and grass have drought legacy effects of approximately 2 years and 1 year, respectively. Statistical analyses partly attribute the differences in drought legacy effects among PFGs to plant eco-hydrological properties (related to traits), including plant water use and hydraulic responses. These results can be used to improve the representation of drought response of different PFGs in land surface models, and assess their biogeochemical and biophysical feedbacks in response to a warmer and drier climate.

Long-term forest resilience to climate change indicated by mortality, regeneration, and growth in semiarid southern Siberia.

Several studies have documented that regional climate warming and the resulting increase in drought stress have triggered increased tree mortality in semiarid forests with unavoidable impacts on regional and global carbon sequestration. Although climate warming is projected to continue into the future, studies examining long-term resilience of semiarid forests against climate change are limited. In this study, long-term forest resilience was defined as the capacity of forest recruitment to compensate for losses from mortality. We observed an obvious change in long-term forest resilience along a local aridity gradient by reconstructing tree growth trend and disturbance history and investigating postdisturbance regeneration in semiarid forests in southern Siberia. In our study, with increased severity of local aridity, forests became vulnerable to drought stress, and regeneration first accelerated and then ceased. Radial growth of trees during 1900-2012 was also relatively stable on the moderately arid site. Furthermore, we found that smaller forest patches always have relatively weaker resilience under the same climatic conditions. Our results imply a relatively higher resilience in arid timberline forest patches than in continuous forests; however, further climate warming and increased drought could possibly cause the disappearance of small forest patches around the arid tree line. This study sheds light on climate change adaptation and provides insight into managing vulnerable semiarid forests.

Trends toward an earlier peak of the growing season in Northern Hemisphere mid-latitudes.

Changes in peak photosynthesis timing (PPT) could substantially change the seasonality of the terrestrial carbon cycle. Spring PPT in dry regions has been documented for some individual plant species on a stand scale, but both the spatio-temporal pattern of shifting PPT on a continental scale and its determinants remain unclear. Here, we use satellite measurements of vegetation greenness to find that the majority of Northern Hemisphere, mid-latitude vegetated area experienced a trend toward earlier PPT during 1982-2012, with significant trends of an average of 0.61 day yr(-1) across 19.4% of areas. These shifts correspond to increased annual accumulation of growing degree days (GDD) due to warming and are most highly concentrated in the eastern United States and Europe. Earlier mean PPT is generally a trait common among areas with summer temperatures higher than 27.6 ± 2.9 °C, summer precipitation lower than 84.2 ± 41.5 mm, and fraction of cold season precipitation greater than 89.2 ± 1.5%. The trends toward earlier PPT discovered here have co-occurred with overall increases in vegetation greenness throughout the growing season, suggesting that summer drought is not a dominant driver of these trends. These results imply that continued warming may facilitate continued shifts toward earlier PPT and cause these trends to become more pervasive, with important implications for terrestrial carbon, water, nutrient, and energy budgets.

Climate-driven speedup of alpine treeline forest growth in the Tianshan Mountains, Northwestern China.

Forest growth is sensitive to interannual climatic change in the alpine treeline ecotone (ATE). Whether the alpine treeline ecotone shares a similar pattern of forest growth with lower elevational closed forest belt (CFB) under changing climate remains unclear. Here, we reported an unprecedented acceleration of Picea schrenkiana forest growth since 1960s in the ATE of Tianshan Mountains, northwestern China by a stand-total sampling along six altitudinal transects with three plots in each transect: one from the ATE between the treeline and the forest line, and the other two from the CFB. All the sampled P. schrenkiana forest patches show a higher growth speed after 1960 and, comparatively, forest growth in the CFB has sped up much slower than that in the ATE. The speedup of forest growth at the ATE is mainly accounted for by climate factors, with increasing temperature suggested to be the primary driver. Stronger water deficit as well as more competition within the CFB might have restricted forest growth there more than that within the ATE, implying biotic factors were also significant for the accelerated forest growth in the ATE, which should be excluded from simulations and predictions of warming-induced treeline dynamics.

Unlocking Drought-Induced Tree Mortality: Physiological Mechanisms to Modeling.

Drought-related tree mortality has become a major concern worldwide due to its pronounced negative impacts on the functioning and sustainability of forest ecosystems. However, our ability to identify the species that are most vulnerable to drought, and to pinpoint the spatial and temporal patterns of mortality events, is still limited. Model is useful tools to capture the dynamics of vegetation at spatiotemporal scales, yet contemporary land surface models (LSMs) are often incapable of predicting the response of vegetation to environmental perturbations with sufficient accuracy, especially under stressful conditions such as drought. Significant progress has been made regarding the physiological mechanisms underpinning plant drought response in the past decade, and plant hydraulic dysfunction has emerged as a key determinant for tree death due to water shortage. The identification of pivotal physiological events and relevant plant traits may facilitate forecasting tree mortality through a mechanistic approach, with improved precision. In this review, we (1) summarize current understanding of physiological mechanisms leading to tree death, (2) describe the functionality of key hydraulic traits that are involved in the process of hydraulic dysfunction, and (3) outline their roles in improving the representation of hydraulic function in LSMs. We urge potential future research on detailed hydraulic processes under drought, pinpointing corresponding functional traits, as well as understanding traits variation across and within species, for a better representation of drought-induced tree mortality in models.

Corrigendum: Unlocking drought-induced tree mortality: Physiological mechanisms to modelling.

[This corrects the article DOI: 10.3389/fpls.2022.835921.].

Modeling vegetation greenness and its climate sensitivity with deep-learning technology.

Climate sensitivity of vegetation has long been explored using statistical or process-based models. However, great uncertainties still remain due to the methodologies' deficiency in capturing the complex interactions between climate and vegetation. Here, we developed global gridded climate-vegetation models based on long short-term memory (LSTM) network, which is a powerful deep-learning algorithm for long-time series modeling, to achieve accurate vegetation monitoring and investigate the complex relationship between climate and vegetation. We selected the normalized difference vegetation index (NDVI) that represents vegetation greenness as model outputs. The climate data (monthly temperature and precipitation) were used as inputs. We trained the networks with data from 1982 to 2003, and the data from 2004 to 2015 were used to validate the models. Error analysis and sensitivity analysis were performed to assess the model errors and investigate the sensitivity of global vegetation to climate change. Results show that models based on deep learning are very effective in simulating and predicting the vegetation greenness dynamics. For models training, the root mean square error (RMSE) is <0.01. Model validation also assure the accuracy of our models. Furthermore, sensitivity analysis of models revealed a spatial pattern of global vegetation to climate, which provides us a new way to investigate the climate sensitivity of vegetation. Our study suggests that it is a good way to integrate deep-learning method to monitor the vegetation change under global change. In the future, we can explore more complex climatic and ecological systems with deep learning and coupling with certain physical process to better understand the nature.

Physiological and environmental control on ecosystem water use efficiency in response to drought across the northern hemisphere.

Drought, a natural hydrometeorological phenomenon, has been more frequent and more widespread due to climate change. Water availability strongly regulates the coupling (or trade-off) between carbon uptake via photosynthesis and water loss through transpiration, known as water-use efficiency (WUE). Understanding the effects of drought on WUE across different vegetation types and along the wet to dry gradient is paramount to achieving better understanding of ecosystem functioning in response to climate change. We explored the physiological and environmental control on ecosystem WUE in response to drought using observations for 44 eddy covariance flux sites in the Northern Hemisphere. We quantified the response of WUE to drought and the relative contributions of gross primary production (GPP) and evapotranspiration (ET) to the variations of WUE. We also examined the control of physiological and environmental factors on monthly WUE under different moisture conditions. Cropland had a peak WUE value under moderate drought conditions, while grassland, deciduous broadleaf forest (DBF), evergreen broadleaf forest (EBF), and evergreen needleleaf forest (ENF) had peak WUE under slight drought conditions. WUE was mainly driven by GPP for cropland, grassland, DBF, and ENF but was mainly driven by ET for EBF. Vapor pressure deficit (VPD) and canopy conductance (Gc) were the most important factors regulating WUE. Moreover, WUE had negative responses to air temperature, precipitation, and VPD but had a positive response to Gc and ecosystem respiration. Our findings highlight the different effects of biotic and abiotic factors on WUE among different vegetation types and the important roles of VPD and Gc in controlling ecosystem WUE in response to drought.

Bedrock geochemistry influences vegetation growth by regulating the regolith water holding capacity.

Although low vegetation productivity has been observed in karst regions, whether and how bedrock geochemistry contributes to the low karstic vegetation productivity remain unclear. In this study, we address this knowledge gap by exploring the importance of bedrock geochemistry on vegetation productivity based on a critical zone investigation across a typical karst region in Southwest China. We show silicon and calcium concentrations in bedrock are strongly correlated with the regolith water loss rate (RWLR), while RWLR can predict vegetation productivity more effectively than previous models. Furthermore, the analysis based on 12 selected karst regions worldwide further suggest that lithological regulation has the potential to obscure and distort the influence of climate change. Our study implies that bedrock geochemistry could exert effects on vegetation growth in karst regions and highlights that the critical role of bedrock geochemistry for the karst region should not be ignored in the earth system model.

Abrupt shift to hotter and drier climate over inner East Asia beyond the tipping point.

Unprecedented heatwave-drought concurrences in the past two decades have been reported over inner East Asia. Tree-ring-based reconstructions of heatwaves and soil moisture for the past 260 years reveal an abrupt shift to hotter and drier climate over this region. Enhanced land-atmosphere coupling, associated with persistent soil moisture deficit, appears to intensify surface warming and anticyclonic circulation anomalies, fueling heatwaves that exacerbate soil drying. Our analysis demonstrates that the magnitude of the warm and dry anomalies compounding in the recent two decades is unprecedented over the quarter of a millennium, and this trend clearly exceeds the natural variability range. The "hockey stick"-like change warns that the warming and drying concurrence is potentially irreversible beyond a tipping point in the East Asian climate system.

Enhanced growth after extreme wetness compensates for post-drought carbon loss in dry forests.

While many studies have reported that drought events have substantial negative legacy effects on forest growth, it remains unclear whether wetness events conversely have positive growth legacy effects. Here, we report pervasive and substantial growth enhancement after extreme wetness by examining tree radial growth at 1929 forest sites, satellite-derived vegetation greenness, and land surface model simulations. Enhanced growth after extreme wetness lasts for 1 to 5 years and compensates for 93 ± 8% of the growth deficit after extreme drought across global water-limited regions. Remarkable wetness-enhanced growths are observed in dry forests and gymnosperms, whereas the enhanced growths after extreme wetness are much smaller in wet forests and angiosperms. Limited or no enhanced growths are simulated by the land surface models after extreme wetness. These findings provide new evidence for improving climate-vegetation models to include the legacy effects of both drought and wet climate extremes.

Does phenology play a role in the feedbacks underlying shrub encroachment?

Shrub encroachment has emerged as a global phenomenon over the past century. Multiple drivers have been put forward to explain the increased shrub dominance in various ecosystems around the world. However, the potential role of phenology in regulating shrub encroachment is not well understood. We address this issue using 3-year continuous monitoring of the phenology of coexisting shrubs and grasses combined with observations of ecohydrological processes (water uptake) and soil conditions (root zone soil moisture, soil texture, and soil temperature) at four study sites in Inner Mongolia, China, with shrub coverage of Caragana microphylla ranging from 0%, to 6.8%, 26.8% and 34.2%. Along such an encroachment gradient, shrubs exhibited progressively earlier onsets and later ends of the growing season, with an overall extension in growing season length by 15 days to 22 days in the later stages of shrub encroachment. Conversely, the coexisting grasses showed earlier occurrences both in spring and autumn phenological phases, which resulted in a phenological gap between shrubs and grasses. Thus, a positive feedback could exist between these phenological changes and shrub encroachment. In shrub patches, soils were wetter, with finer texture, and with more suitable temperatures for plant survival and development, which favored the lengthening of growing season of shrubs. The longer growing seasons are associated with longer periods of water use and photosynthesis for shrubs, and better opportunities for water uptake, with the overall effect of facilitating shrub growth and further expansion.

Responses of soil respiration to rainfall addition in a desert ecosystem: Linking physiological activities and rainfall pattern.

The tight linkage between photosynthesis (An) and soil respiration (Rs) has been verified in many terrestrial ecosystems. However, it remains unclear whether this linkage occurs in desert ecosystems, where water is considered an important trigger of carbon cycling. A field experiment was performed under seven simulated rainfall amounts (0, 3, 5, 10, 15, 25, and 40 mm) with two co-existing desert plants (Reaumuria soongorica and Nitraria sphaerocarpa) in June (early growing season, EGS) and August (middle growing season, MGS) in 2016. An, Rs, predawn water potential (Ψpd), soil temperature (Ts) and soil moisture (Swc) were measured for each treatment or control plot for 3 weeks. Our objective was to examine the effects of rainfall pattern on Rs and physiological responses of the two plants and the relationships between Rs and biotic and abiotic factors. No obvious variations in Ψpd or An were found under small rainfall events. However, when the rainfall amount exceeded 10 mm, both plants responded strongly, and the response patterns of Rs showed trends similar to those of An, which varied between species and seasons. Moreover, rain additions of 3-40 mm significantly increased Rs, and the relative changes in Rs (ΔRs) of both species were much larger in the EGS than in the MGS. Importantly, abiotic factors may have controlled the variations in Rs under small rain events while An played a more important role in regulating the variations in Rs when the rainfall amount exceeded 10 mm for both species, suggest that the rainfall pattern-driven changes in Rs composition interact with physiological activity and abiotic factors to regulate the response of Rs to rainfall variability in desert ecosystems. Thus, climate change in the coming decades may lead to carbon sequestration by desert plants, which may cause desert ecosystems to act as carbon sinks.

Exposures to temperature beyond threshold disproportionately reduce vegetation growth in the northern hemisphere.

In recent decades, terrestrial vegetation in the northern hemisphere (NH) has been exposed to warming and more extremely high temperatures. However, the consequences of these changes for terrestrial vegetation growth remain poorly quantified and understood. By examining a satellite-based vegetation index, tree-ring measurements and land-surface model simulations, we discovered a consistent convex pattern in the responses of vegetation growth to temperature exposure (TE) for forest, shrub and grass in both the temperate (30°-50° N) and boreal (50°-70° N) NH during the period of 1982-2012. The response of vegetation growth to TE for the three vegetation types in both the temperate and boreal NH increased convergently with increasing temperature, until vegetation type-dependent temperature thresholds were reached. A TE beyond these temperature thresholds resulted in disproportionately weak positive or even strong negative responses. Vegetation growth in the boreal NH was more vulnerable to extremely high-temperature events than vegetation growth in the temporal NH. The non-linear responses discovered here provide new insights into the dynamics of northern terrestrial ecosystems in a warmer world.

Phytoliths and phytolith carbon occlusion in aboveground vegetation of sandy grasslands in eastern Inner Mongolia, China.

Grasslands play a crucial role in the coupled biogeochemical cycles of carbon (C) and silicon (Si) because they have a large biogenic Si pool (i.e. phytoliths). In recent decades, desertification has occurred extensively in sandy grasslands due to human activities and to increased aridity as a consequence of climate change. The present study determined the contents of phytoliths and C occlusion within phytoliths (PhytOC) in sandy grassland with different vegetation coverage from eastern Inner Mongolia, China and preliminarily assessed the effects of desertification on phytoliths and PhytOC production. Our results showed that the phytolith and PhytOC contents among different plant species varied from 0.68 to 9.23% and 0.03 to 1.13‰, respectively. However, the community-weighted means of the phytolith and PhytOC contents for the total aboveground vegetation were only 1.13-3.61% and 0.09-0.35‰, respectively, and their respective production fluxes ranged from 8.94 to 47.8 kg ha-1 year-1 and from 0.06 to 0.48 kg ha-1 year-1, respectively. As desertification progressed, the total contents of phytoliths and PhytOC in aboveground vegetation did not change significantly, whereas the production fluxes of phytoliths and PhytOC were markedly reduced. This study indicates that grassland desertification decreases the range of the total contents of phytolith and PhytOC by reducing species richness, and decreases the production fluxes of phytoliths and PhytOC by reducing aboveground biomass. Grassland restoration can theoretically enhance the production fluxes of phytoliths and PhytOC ~ five-fold.