Simulated climate change decreases nutrient resorption from senescing leaves

Nutrient resorption is the process whereby plants recover nutrients from senescing leaves and reallocate them to storage structures or newer tissues. Elemental resorption of foliar N and P has been shown to respond to temperature and precipitation, but we know remarkably little about the influence of warming and drought on the resorption of these and other essential plant macro‐ and micronutrients, which could alter the ability of species to recycle their nutrients. We conducted a 5 year manipulative field study to simulate predicted climate change conditions and studied the effects of warming (W), rainfall reduction (RR), and their combination (W+RR) on nutrient resorption efficiency in five coexisting shrub species in a semiarid shrubland. Both mature and senesced leaves showed significant reductions in their nutrient contents and an altered stoichiometry in response to climate change conditions. Warming (W, W+RR) reduced mature leaf N, K, Ca, S, Fe, and Zn and senesced leaf N, Ca, Mg, S, Fe, and Zn contents relative to ambient temperature conditions. Warming increased mature leaf C/N ratios and decreased N/P and C/P ratios and increased senesced leaf C/N and C/P ratios. Furthermore, W and W+RR reduced nutrient resorption efficiencies for N (6.3%), K (19.8%), S (70.9%) and increased Ca and Fe accumulation in senesced leaves (440% and 35.7%, respectively) relative to the control treatment. Rainfall reduction decreased the resorption efficiencies of N (6.7%), S (51%), and Zn (46%). Reductions in nutrient resorption efficiencies with warming and/or rainfall reduction were rather uniform and consistent across species. The negative impacts of warming and rainfall reduction on foliar nutrient resorption efficiency will likely cause an impairment of plant nutrient budgets and fitness across coexisting native shrubs in this nutrient‐poor habitat, with probable implications for key ecosystem functions such as reductions in nutrient retention in vegetation, litter decomposition, and nutrient cycling rates.     

Alteration of the phenology of leaf senescence and fall in winter deciduous species by climate change: effects on nutrient proficiency

Leaf senescence in winter deciduous species signals the transition from the active to the dormant stage. The purpose of leaf senescence is the recovery of nutrients before the leaves fall. Photoperiod and temperature are the main cues controlling leaf senescence in winter deciduous species, with water stress imposing an additional influence. Photoperiod exerts a strict control on leaf senescence at latitudes where winters are severe and temperature gains importance in the regulation as winters become less severe. On average, climatic warming will delay and drought will advance leaf senescence, but at varying degrees depending on the species. Warming and drought thus have opposite effects on the phenology of leaf senescence, and the impact of climate change will therefore depend on the relative importance of each factor in specific regions. Warming is not expected to have a strong impact on nutrient proficiency although a slower speed of leaf senescence induced by warming could facilitate a more efficient nutrient resorption. Nutrient resorption is less efficient when the leaves senesce prematurely as a consequence of water stress. The overall effects of climate change on nutrient resorption will depend on the contrasting effects of warming and drought. Changes in nutrient resorption and proficiency will impact production in the following year, at least in early spring, because the construction of new foliage relies almost exclusively on nutrients resorbed from foliage during the preceding leaf fall. Changes in the phenology of leaf senescence will thus impact carbon uptake, but also ecosystem nutrient cycling, especially if the changes are consequence of water stress.     

Environmental and physiological factors governing nutrient resorption efficiency in barley

Summary Barley plants (Hordeum vulgare cv. stepto) were grown in a greenhouse under two nutrient and water levels and four treatments intended to alter sourcesink relationships, in a factorial experiment designed to study factors governing efficiency of nutrient resorption from senescing leaves. Plant growth was enhanced in high-nutrient treatments, leading to higher concentrations of nitrogen (N) and phosphorus (P) in leaves and ears. Water stress reduced growth, but plants in waterstressed treatments had equal or higher nutrient concentrations than watered plants. Nutrient resorption efficiency was higher at low than at high nutrient availability, and was higher in watered than in water-stressed plants. Treatments in which sink strength was increased had enhanced resorption efficiency, as well as those in which the source activity was reduced. Our data show that the amount of nutrient resorbed and the efficiency of the resorption process depend on plant nutrient and water status, and that the presence of an active sink strongly enhances nutrient resorption.     

Climate change has only a minor impact on nutrient resorption parameters in a high-latitude peatland

Nutrient resorption from senescing plant tissues is an important determinant of the fitness of plant populations in nutrient-poor ecosystems, because it makes plants less dependent on current nutrient uptake. Moreover, it can have significant “afterlife” effects through its impact on litter chemistry and litter decomposability. Little is known about the effects of climate change on nutrient resorption. We studied the effects of climate change treatments (including winter snow addition, and spring and/or summer warming) on nutrient resorption of four dominant species in a nutrient-poor subarctic peatland. These species were Betula nana (woody deciduous), Vaccinium uliginosum (woody deciduous), Calamagrostis lapponica (graminoid) and Rubus chamaemorus (forb). After five years of treatments both mature and senesced leaf N concentrations showed a small but significant overall reduction in response to the climate treatments. However, the effects were species-specific. For example, in the controls the N concentration in senesced leaves of Calamagrostis (3.0±0.2 mg N g⁻¹) was about four times lower than for Rubus (11.2±0.2 mg N g⁻¹). There were no significant treatment effects on N resorption efficiency (% of the N pool in mature leaves that is resorbed during senescence). The nitrogen resorption efficiency of Calamagrostis (about 80%) was higher than in the other three species (about 60%). Thus, climate change has only a minor impact on nutrient resorption parameters. However, given the substantial interspecific differences in these parameters, substantial changes in plant–soil feedbacks may be expected as a result of the observed changes in the species composition of high-latitude vegetation. These changes are species-specific and thus difficult to predict.     

Lasting effects of climate disturbance on perennial grassland above‐ground biomass production under two cutting frequencies

Climate extremes can ultimately reshape grassland services such as forage production and change plant functional type composition. This 3‐year field research studied resistance to dehydration and recovery after rehydration of plant community and plant functional types in an upland perennial grassland subjected to climate and cutting frequency (Cut+, Cut?) disturbances by measuring green tissue percentage and above‐ground biomass production (ANPP). In year 1, a climate disturbance gradient was applied by co‐manipulating temperature and precipitation. Four treatments were considered: control and warming‐drought climatic treatment, with or without extreme summer event. In year 2, control and warming‐drought treatments were maintained without extreme. In year 3, all treatments received ambient climatic conditions. We found that the grassland community was very sensitive to dehydration during the summer extreme: aerial senescence reached 80% when cumulated climatic water balance fell to ?156 mm and biomass declined by 78% at the end of summer. In autumn, canopy greenness and biomass totally recovered in control but not in the warming‐drought treatment. However ANPP decreased under both climatic treatments, but the effect was stronger on Cut+ (?24%) than Cut? (?15%). This decline was not compensated by the presence of three functional types because they were negatively affected by the climatic treatments, suggesting an absence of buffering effect on grassland production. In the following 2 years, lasting effects of climate disturbance on ANPP were observable. The unexpected stressful conditions of year 3 induced a decline in grassland production in the Cut+ control treatment. The fact that this treatment cumulated higher (45%) N export over the 3 years suggests that N plays a key role in ANPP stability. As ANPP in this mesic perennial grassland did not show engineering resilience, long‐term experimental manipulation is needed. Infrequent mowing appears more appropriate for sustaining grassland ANPP under future climate extremes.     

Resistance of soil protein depolymerization rates to eight years of elevated CO<Subscript>2</Subscript>, warming,and summer drought in a temperate heathland

Soil N availability for plants and microorganisms depends on the breakdown of soil polymers such as proteins into smaller, assimilable units by microbial extracellular enzymes. Changing climatic conditions are expected to alter protein depolymerization rates over the next decades, and thereby affect the potential for plant productivity. We here tested the effect of increased CO₂ concentration, temperature, and drought frequency on gross rates of protein depolymerization, N mineralization, microbial amino acid and ammonium uptake using ¹⁵N pool dilution assays. Soils were sampled in fall 2013 from the multifactorial climate change experiment CLIMAITE that simulates increased CO₂ concentration, temperature, and drought frequency in a fully factorial design in a temperate heathland. Eight years after treatment initiation, we found no significant effect of any climate manipulation treatment, alone or in combination, on protein depolymerization rates. Nitrogen mineralization, amino acid and ammonium uptake showed no significant individual treatment effects, but significant interactive effects of warming and drought. Combined effects of all three treatments were not significant for any of the measured parameters. Our findings therefore do not suggest an accelerated release of amino acids from soil proteins in a future climate at this site that could sustain higher plant productivity.     

Modest enhancement of nitrogen conservation via retranslocation in response to gradients in N supply and leaf N status

Plant nutrient resorption, a ubiquitous mechanism of nutrient conservation, has often been proposed to be more pronounced in infertile than fertile habitats, and in species common to infertile compared to fertile habitats, because of the presumed advantage when nutrients are scarce. However, previous studies provide weak and inconsistent empirical support for these hypotheses, although few have examined intraspecific variation across well-quantified resource gradients. This study addresses intraspecific patterns of nutrient resorption for eight species across two N availability gradients on similar soils in an N-limited oak savanna ecosystem: a long-term fire frequency gradient with a negatively correlated N fertility gradient and a long-term N fertilization gradient. We hypothesized that both resorption proficiency (the minimum nutrient level retained in a senesced leaf) and efficiency (the proportional change in leaf nutrient concentration) would decrease with increasing soil N availability and plant N status. For the seven non-N fixers, either resorption proficiency or efficiency decreased modestly in treatments with higher N availability. In contrast, the legume Amorpha canescens Pursh had higher N levels in green and senesced leaves, and resorbed N much more weakly than the non-fixers, and did not respond in terms of proficiency or efficiency to soil N availability. Across all species and sites in each N fertility gradient, a scaling analysis showed greater resorption efficiency in plants with lower N concentrations. Our data suggest that species can have modest resorption responses reflective of soil nutrient availability and differences in resorption related to their N economy that represent mechanisms of nutrient conservation in nutrient-limited soils.     

Warming and drought reduce temperature sensitivity of nitrogen transformations

Shifts in nitrogen (N) mineralization and nitrification rates due to global changes can influence nutrient availability, which can affect terrestrial productivity and climate change feedbacks. While many single‐factor studies have examined the effects of environmental changes on N mineralization and nitrification, few have examined these effects in a multifactor context or recorded how these effects vary seasonally. In an old‐field ecosystem in Massachusetts, USA, we investigated the combined effects of four levels of warming (up to 4 °C) and three levels of precipitation (drought, ambient, and wet) on net N mineralization, net nitrification, and potential nitrification. We also examined the treatment effects on the temperature sensitivity of net N mineralization and net nitrification and on the ratio of C mineralization to net N mineralization. During winter, freeze–thaw events, snow depth, and soil freezing depth explained little of the variation in net nitrification and N mineralization rates among treatments. During two years of treatments, warming and altered precipitation rarely influenced the rates of N cycling, and there was no evidence of a seasonal pattern in the responses. In contrast, warming and drought dramatically decreased the apparent Q₁₀ of net N mineralization and net nitrification, and the warming‐induced decrease in apparent Q₁₀ was more pronounced in ambient and wet treatments than the drought treatment. The ratio of C mineralization to net N mineralization varied over time and was sensitive to the interactive effects of warming and altered precipitation. Although many studies have found that warming tends to accelerate N cycling, our results suggest that warming can have little to no effect on N cycling in some ecosystems. Thus, ecosystem models that assume that warming will consistently increase N mineralization rates and inputs of plant‐available N may overestimate the increase in terrestrial productivity and the magnitude of an important negative feedback to climate change.     

Resorption proficiency and efficiency of leaf nutrients in woody plants in eastern China

Aims (i) To explore variations in nutrient resorption of woody plants and their relationship with nutrient limitation and (ii) to identify the factors that control these variations in forests of eastern China.Methods We measured nitrogen (N) and phosphorus (P) concentrations in both green and senesced leaves of 172 woody species at 10 forest sites across eastern China. We compared the nutrient resorption proficiency (NuRP) and efficiency (NuRE) of N and P in plant leaves for different functional groups; we further investigated the latitudinal and altitudinal variations in NuRP and NuRE and the impacts of climate, soil and plant types on leaf nutrient resorptions.Important findings On average, the leaf N resorption proficiency (NRP) and P resorption proficiency (PRP) of woody plants in eastern China were 11.1mg g ? 1 and 0.65 mg g ? 1, respectively; and the corresponding N resorption efficiency (NRE) and P resorption efficiency (PRE) were 49.1% and 51.0%, respectively. Angiosperms have higher NRP (are less proficient) values and lower NRE and PRE values than gymnosperms, but there are no significant differences in NRP, PRP and PRE values between species with different leaf habits (evergreen vs. deciduous angiosperms). Trees have higher NRE and PRE than shrubs. Significant geographical patterns of plant nutrient resorption exist in forests of eastern China. In general, NRP and PRE decrease and PRP and NRE increase with increasing latitude/altitude for all woody species and for the different plant groups. Plant functional groups show more controls than environmental factors (climate and soil) on the N resorption traits (NRP and NRE), while site-related variables present more controls than plant types on PRP and PRE. NRP increases and PRP and NRE decrease significantly with increasing temperature and precipitation for the overall plants and for most groups, except that significant PRE–climate relationship holds for only evergreen angiosperms. Leaf nutrient resorption did not show consistent responses in relation to soil total N and P stoichiometry, probably because the resorption process is regulated by the relative costs of drawing nutrients from soil versus from senescing leaves. These results support our hypothesis that plants growing in P-limited habitats (low latitudes/altitudes or areas with high precipitation/temperature) should have lower PRP and higher PRE, compared with their counterparts in relatively N-limited places (high latitudes/altitudes or areas with low precipitation/temperature). Our findings can improve the understanding of variations in N and P resorption and their responses to global change, and thus facilitate to incorporate these nutrient resorption processes into future biogeochemical models.     

Reproductive output in Mediterranean shrubs under climate change experimentally induced by drought and warming

The effects of climate change on plant reproductive performance affects the sequence of different plant reproductive stages from flowering to seed production and viability, as well as the network of relationships between them. These effects are expected to respond to different components of climate change, such as temperature and water availability, and may be sensitive to differences in species phenology.We used long-term experimental drought and warming treatments to study the effect of climate change on flower production, fruit and seed-set, seed size and seed germination rate (proportion of germinating seeds) in three Mediterranean shrubs coexisting in a coastal shrubland.Larger plants produced significantly more flowers in all three species, and higher fruit-set in Dorycnium pentaphyllum. Flower production was reduced in drought and warming treatments in the spring-flowering species D. pentaphyllum and Helianthemum syriacum, but not in the autumn–winter species Erica multiflora, which increased flowering in the warming treatment. However, the drought treatment eventually resulted in a decreased seed-set in E. multiflora. Structural equation modelling revealed strong correlations between the sequential reproductive stages. Specifically, flower density in inflorescences determined seed-set in H. syriacum, and seed size and germination rate in E. multiflora. Nevertheless, the relevance of relationships between reproductive traits changed between climatic treatments: in D. pentaphyllum a direct relationship between plant size and seed size only arised in the drought treatment, while in H. syriacum climate treatments resulted in a stronger relationship between the number of flowers and seed-set.This experimental study shows the ability of changing climatic variables to determine the reproductive sequential process of woody species. We show that several parameters of the reproductive performance of some Mediterranean species are affected by drought and warming treatments simulating climate change, highlighting the importance of changes in both water availability and temperature, and the sequential relationship between reproductive stages. Phenological patterns also contribute to species’ differential responses to climatic change, due to the relationship of these patterns with resource availability, environmental conditions and plant–pollinator interactions.     

不同降水条件下荒漠草原植物的养分含量及回收对增温和氮素添加的响应

为了探讨荒漠草原植物养分回收特征对长期增温和氮素添加的响应以及自然降水变异对其的调控作用,该研究依托实施12年的模拟增温和氮素添加实验平台,在相对多雨的2016年(超过长期均值52%)和相对少雨的2017年(低于长期均值16%),以常见C_3植物银灰旋花(Convolvulus ammannii)和C_4植物木地肤(Kochia prostrata)为研究对象,测定分析绿叶和枯叶的氮磷含量及回收效率。结果表明:(1)在相对多雨年(2016年),增温使2种植物的绿叶氮、枯叶氮、绿叶磷、枯叶磷含量分别增加了14.32%、25.45%、17.97%和46.47%,氮、磷回收效率分别显著减小了9.41%和16.99%(P0.05);氮素添加使2种植物的绿叶氮、枯叶氮、绿叶磷、枯叶磷含量分别提高了17.32%、25.62%、20.21%和51.41%,而氮、磷回收效率显著降低了9.33%和18.89%(P0.05);增温+氮素添加共同处理显著增加了植物氮磷含量、降低了氮磷回收效率。(2)在相对少雨年(2017年),增温、氮素添加、增温+氮素添加处理对植物叶片氮磷含量、回收效率均无显著影响。(3)叶片氮磷含量在物种间差异极显著(P0.000 1),而氮磷回收效率在物种间无显著差异。(4)回归分析表明,植物叶片氮磷含量随着土壤无机氮、有效磷及含水量的增加而增加,植物氮磷回收效率则随着土壤养分和水分的可利用性的增加而降低。研究认为,荒漠草原植物养分回收对全球变化的响应受自然降水变异的调控。     

Interplay between abiotic (drought) and biotic (virus) stresses in tomato plants

With climate warming, drought becomes a vital challenge for agriculture. Extended drought periods affect plant–pathogen interactions. We demonstrate an interplay in tomato between drought and infection with tomato yellow leaf curl virus (TYLCV). Infected plants became more tolerant to drought, showing plant readiness to water scarcity by reducing metabolic activity in leaves and increasing it in roots. Reallocation of osmolytes, such as carbohydrates and amino acids, from shoots to roots suggested a role of roots in protecting infected tomatoes against drought. To avoid an acute response possibly lethal for the host organism, TYLCV down-regulated the drought-induced activation of stress response proteins and metabolites. Simultaneously, TYLCV promoted the stabilization of osmoprotectants' patterns and water balance parameters, resulting in the development of buffering conditions in infected plants subjected to prolonged stress. Drought-dependent decline of TYLCV amounts was correlated with HSFA1-controlled activation of autophagy, mostly in the roots. The tomato response to combined drought and TYLCV infection points to a mutual interaction between the plant host and its viral pathogen.     

Mapping nutrient resorption efficiencies of subarctic cryptogams and seed plants onto the Tree of Life

Nutrient resorption from senescing photosynthetic organs is a powerful mechanism for conserving nitrogen (N) and phosphorus (P) in infertile environments. Evolution has resulted in enhanced differentiation of conducting tissues to facilitate transport of photosynthate to other plant parts, ultimately leading to phloem. Such tissues may also serve to translocate N and P to other plant parts upon their senescence. Therefore, we hypothesize that nutrient resorption efficiency (RE, % of nutrient pool exported) should correspond with the degree of specialization of these conducting tissues across the autotrophic branches of the Tree of Life. To test this hypothesis, we had to compare members of different plant clades and lichens within a climatic region, to minimize confounding effects of climatic drivers on nutrient resorption. Thus, we compared RE among wide‐ranging basal clades from the principally N‐limited subarctic region, employing a novel method to correct for mass loss during senescence. Even with the limited numbers of species available for certain clades in this region, we found some consistent patterns. Mosses, lichens, and lycophytes generally showed low RE_N (<20%), liverworts and conifers intermediate (40%) and monilophytes, eudicots, and monocots high (>70%). RE_P appeared higher in eudicots and liverworts than in mosses. Within mosses, taxa with more efficient conductance also showed higher RE_N. The differences in RE_N among clades broadly matched the degree of specialization of conducting tissues. This novel mapping of a physiological process onto the Tree of Life broadly supports the idea that the evolution of conducting tissues toward specialized phloem has aided land plants to optimize their internal nitrogen recycling. The generality of evolutionary lines in conducting tissues and nutrient resorption efficiency needs to be tested across different floras in different climatic regions with different levels of N versus P availability.     

Nutrient resorption patterns of plant functional groups in a tropical savanna: variation and functional significance

Green and senesced leaf nitrogen (N) and phosphorus (P) concentrations of different plant functional groups in savanna communities of Kruger National Park, South Africa were analyzed to determine if nutrient resorption was regulated by plant nutritional status and foliar N:P ratios. The N and P concentrations in green leaves and the N concentrations in senesced leaves differed significantly between the dominant plant functional groups in these savannas: fine-leaved trees, broad-leaved trees and grasses. However, all three functional groups reduced P to comparable and very low levels in senesced leaves, suggesting that P was tightly conserved in this tropical semi-arid savanna ecosystem. Across all functional groups, there was evidence for nutritional control of resorption in this system, with both N and P resorption efficiencies decreasing as green leaf nutrient concentrations increased. However, specific patterns of resorption and the functional relationships between nutrient concentrations in green and senesced leaves varied by nutrient and plant functional group. Functional relationships between N concentrations in green and senesced leaves were indistinguishable between the dominant groups, suggesting that variation in N resorption efficiency was largely the result of inter-life form differences in green leaf N concentrations. In contrast, observed differences in P resorption efficiencies between life forms appear to be the result of both differences in green leaf P concentrations as well as inherent differences between life forms in the fraction of green leaf P resorbed from senescing leaves. Our results indicate that foliar N:P ratios are poor predictors of resorption efficiency in this ecosystem, in contrast to N and P resorption proficiencies, which are more responsive to foliar N:P ratios.     

Interactive responses of old‐field plant growth and composition to warming and precipitation

As Earth's atmosphere accumulates carbon dioxide (CO₂) and other greenhouse gases, Earth's climate is expected to warm and precipitation patterns will likely change. The manner in which terrestrial ecosystems respond to climatic changes will in turn affect the rate of climate change. Here we describe responses of an old‐field herbaceous community to a factorial combination of four levels of warming (up to 4 °C) and three precipitation regimes (drought, ambient and rain addition) over 2 years. Warming suppressed total production, shoot production, and species richness, but only in the drought treatment. Root production did not respond to warming, but drought stimulated the growth of deeper (> 10 cm) roots by 121% in 1 year. Warming and precipitation treatments both affected functional group composition, with C₄ grasses and other annual and biennial species entering the C₃ perennial‐dominated community in ambient rainfall and rain addition treatments as well as in warmed treatments. Our results suggest that, in this mesic system, expected changes in temperature or large changes in precipitation alone can alter functional composition, but they have little effect on total herbaceous plant growth. However, drought limits the capacity of the entire system to withstand warming. The relative insensitivity of our study system to climate suggests that the herbaceous component of old‐field communities will not dramatically increase production in response to warming or precipitation change, and so it is unlikely to provide either substantial increases in forage production or a meaningful negative feedback to climate change later this century.     

Combined effects of nitrogen addition and litter manipulation on nutrient resorption of Leymus chinensis in a semi‐arid grassland of northern China

Plant growth in semi‐arid ecosystems is usually severely limited by soil nutrient availability. Alleviation of these resource stresses by fertiliser application and aboveground litter input may affect plant internal nutrient cycling in such regions. We conducted a 4‐year field experiment to investigate the effects of nitrogen (N) addition (10 g N·m^?2·year^?1) and plant litter manipulation on nutrient resorption of Leymus chinensis, the dominant native grass in a semi‐arid grassland in northern China. Although N addition had no clear effects on N and phosphorus (P) resorption efficiencies in leaves and culms, N fertilisation generally decreased leaf N resorption proficiency by 54%, culm N resorption proficiency by 65%. Moreover, N fertilisation increased leaf P resorption proficiency by 13%, culm P resorption proficiency by 20%. Under ambient or enriched N conditions, litter addition reduced N and P resorption proficiencies in both leaves and culms. The response of P resorption proficiency to litter manipulation was more sensitive than N resorption proficiency: P resorption proficiency in leaves and culms decreased strongly with increasing litter amount under both ambient and enriched N conditions. In contrast, N resorption proficiency was not significantly affected by litter addition, except for leaf N resorption proficiency under ambient N conditions. Furthermore, although litter addition caused a general decrease of leaf and culm nutrient resorption efficiencies under both ambient and enriched N conditions, litter addition effects on nutrient resorption efficiency were much weaker than the effects of litter addition on nutrient resorption proficiency. Taken together, our results show that leaf and non‐leaf organs of L. chinensis respond consistently to altered soil N availability. Our study confirms the strong effects of N addition on plant nutrient resorption processes and the potential role of aboveground litter, the most important natural fertiliser in terrestrial ecosystems, in influencing plant internal nutrient cycling.     

Net nitrogen mineralization and leaching in response to warming and nitrogen deposition in a temperate old field: the importance of winter temperature

While climate warming can increase plant N availability over the growing season by increasing rates of N mineralization, increased N mineralization over winter at a time when plant roots are largely inactive, coupled with an increased frequency of soil freeze–thaw cycles, may increase soil N leaching losses. We examined changes in soil net N mineralization and N leaching in response to warming and N addition (6 g m^?2 year^?1) in a factorial experiment conducted in a temperate old field. We used two warming treatments, year-round and winter-only warming, to isolate the effects of winter warming on soil N dynamics from the year-round warming effects. We estimated net N mineralization using in situ soil cores with resin bags placed at the bottom to catch throughput, and we measured N leaching using lysimeters located below the plant rooting zone at a depth of 50 cm. There were minor effects of warming on changes in soil extractable N and resin N in the soil cores over winter. Nevertheless, the overall effects of both warming and N addition on net N mineralization (the sum of changes in soil extractable N and resin N) were not significant over this period. Likewise, there were no significant treatment effects on the concentration of N in leachate collected below the plant rooting zone. However, in response to winter warming, net N mineralization over summer was approximately double that of both the ambient and year-round warming treatments. This result demonstrates a potentially large and unexpected effect of winter warming on soil N availability in this old field system.     

Drought-deciduous behavior reduces nutrient losses from temperate deciduous trees under severe drought

Nutrient resorption from senescing leaves is an important mechanism of nutrient conservation in temperate deciduous forests. Resorption, however, may be curtailed by climatic events that cause rapid leaf death, such as severe drought, which has been projected to double by the year 2100 in the eastern United States. During a record drought in the southeastern US, we studied 18 common temperate winter-deciduous trees and shrubs to understand how extreme drought affects nutrient resorption of the macronutrients N, P, K, and Ca. Four species exhibited drought-induced leaf senescence and maintained higher leaf water potentials than the remaining 14 species (here called drought-evergreen species). This strategy prevented extensive leaf desiccation during the drought and successfully averted large nutrient losses caused by leaf desiccation. These four drought-deciduous species were also able to resorb N, P, and K from drought-senesced leaves, whereas drought-evergreen species did not resorb any nutrients from leaves lost to desiccation during the drought. For Oxydendrum arboreum, the species most severely affected by the drought, our results indicate that trees lost 50% more N and P due to desiccation than would have been lost from fall senescence alone. For all drought-deciduous species, resorption of N and P in fall-senesced leaves was highly proficient, whereas resorption was incomplete for drought-evergreen species. The lower seasonal nutrient losses of drought-deciduous species may give them a competitive advantage over drought-evergreen species in the years following the drought, thereby impacting species composition in temperate deciduous forests in the future.     

Photosynthesis and growth reduction with warming are driven by nonstomatal limitations in a Mediterranean semi‐arid shrub

Whereas warming enhances plant nutrient status and photosynthesis in most terrestrial ecosystems, dryland vegetation is vulnerable to the likely increases in evapotranspiration and reductions in soil moisture caused by elevated temperatures. Any warming‐induced declines in plant primary production and cover in drylands would increase erosion, land degradation, and desertification. We conducted a four‐year manipulative experiment in a semi‐arid Mediterranean ecosystem to evaluate the impacts of a ~2°C warming on the photosynthesis, transpiration, leaf nutrient status, chlorophyll content, isotopic composition, biomass growth, and postsummer survival of the native shrub Helianthemum squamatum. We predicted that warmed plants would show reduced photosynthetic activity and growth, primarily due to the greater stomatal limitation imposed by faster and more severe soil drying under warming. On average, warming reduced net photosynthetic rates by 36% across the study period. Despite this strong response, warming did not affect stomatal conductance and transpiration. The reduction of peak photosynthetic rates with warming was more pronounced in a drought year than in years with near‐average rainfall (75% and 25–40% reductions relative to controls, respectively), with no indications of photosynthetic acclimation to warming through time. Warmed plants had lower leaf N and P contents, δ¹³C, and sparser and smaller leaves than control plants. Warming reduced shoot dry mass production by 31%. However, warmed plants were able to cope with large reductions in net photosynthesis, leaf area, and shoot biomass production without changes in postsummer survival rates. Our findings highlight the key role of nonstomatal factors (biochemical and/or nutritional) in reducing net carbon assimilation rates and growth under warming, which has important implications for projections of plant carbon balance under the warmer and drier climatic scenario predicted for drylands worldwide. Projected climate warming over the coming decades could reduce net primary production by about one‐third in semi‐arid gypsum shrublands dominated by H. squamatum.     

Patterns and Mechanisms of Nutrient Resorption in Plants

Nutrient resorption (NR) plays a key role in the nutrient conservation of plants. However, a fundamental understanding of the mechanisms that control NR remains limited. In this review, we examine how intrinsic controls (e.g., genetic variability and plant development) and extrinsic environmental controls (e.g., climate and soil fertility) influence NR. We also examined conceptual NR advances, mass loss correction, measurement in non-leaf plant tissues for whole-plant nutrient budget accounting, and the use of stoichiometric ratios in place of individual elements. Nutrient resorption from senescing leaves is greater than that from stems/culms or roots. Nutrients resorbed from stems and roots in woody plants are lower than in non-woody plants. Deciduous plants are more efficient in resorbing leaf nutrients prior to senescence than are evergreen plants. Furthermore, reproductive efforts tend to increase NR. Along a latitudinal gradient of terrestrial biomes, nitrogen resorption efficiency decreases and phosphorus resorption efficiency increases with increasing temperature and precipitation; however, latitudinal patterns reflect the influences of several coupling factors such as genetic variation, climate, soil, and disturbance history. Nutrient fertilization experiments have demonstrated that increased soil fertility reduces NR. The inquiries into the impacts of ongoing climate change on NR are still at a nascent stage. Future NR studies are needed to better understand the independent effects of a wide range of genetic variation, plant development, and environment, and possibly the different responses of plants to environmental change; particularly elevated atmospheric CO₂ concentrations and global warming.