Climate gradient and leaf carbon investment influence the effects of climate change on water use efficiency of forests: A meta-analysis.

Forest ecosystems cover a large area of the global land surface and are important carbon sinks. The water-carbon cycles of forests are prone to climate change, but uncertainties remain regarding the magnitude of water use efficiency (WUE) response to climate change and the underpinning mechanism driving WUE variation. We conducted a meta-analysis of the effects of elevated CO2 concentration (eCO2 ), drought and elevated temperature (eT) on the leaf- to plant-level WUE, covering 80 field studies and 95 tree species. The results showed that eCO2 increased leaf intrinsic and instantaneous WUE (WUEi, WUEt), whereas drought enhanced both leaf- and plant-level WUEs. eT increased WUEi but decreased carbon isotope-based WUE, possibly due to the influence of mesophyll conductance. Stimulated leaf-level WUE by drought showed a progressing trend with increasing latitude, while eCO2 -induced WUE enhancement showed decreasing trends after >40° N. These latitudinal gradients might influence the spatial pattern of climate and further drove WUE variation. Moreover, high leaf-level WUE under eCO2 and drought was accompanied by low leaf carbon contents. Such a trade-off between growth efficiency and defence suggests a potentially compromised tolerance to diseases and pests. These findings add important ecophysiological parameters into climate models to predict carbon-water cycles of forests.

Leaf water relations determine the trade-off between ozone resistance and stomatal functionality in urban tree species.

Urban trees possess different capacities to mitigate ozone (O3) pollution through stomatal uptake. Stomatal closure protects trees from oxidative damage but limits their growth. To date, it is unclear how plant hydraulic function affect stomatal behaviour and determine O3 resistance. We assessed gas exchange and hydraulic traits in three subtropical urban tree species, Celtis sinensis, Quercus acutissima, and Q. nuttallii, under nonfiltered ambient air (NF) and elevated O3 (NF60). NF60 decreased photosynthetic rate (An) and stomatal conductance (gs) only in Q. acutissima and Q. nuttallii. Maintained An in C. sinensis suggested high O3 resistance and was attributed to higher leaf capacitance at the full turgor. However, this species exhibited a reduced stomatal sensitivity to vapour pressure deficit and an increased minimal gs under NF60. Such stomatal dysfunction did not decrease intrinsic water use efficiency (WUE) due to a tight coupling of An and gs. Conversely, Q. acutissima and Q. nuttallii showed maintained stomatal sensitivity and increased WUE, primarily correlated with gs and leaf water relations, including relative water content and osmotic potential at turgor loss point. Our findings highlight a trade-off between O3 resistance and stomatal functionality, with efficient stomatal control reducing the risk of hydraulic failure under combined stresses.

Drought mitigates the adverse effects of O3 on plant photosynthesis rather than growth: A global meta-analysis considering plant functional types.

Tropospheric ozone (O3 ) is a phytotoxic air pollutant adversely affecting plant growth. High O3 exposures are often concurrent with summer drought. The effects of both stresses on plants are complex, and their interactions are not yet well understood. Here, we investigate whether drought can mitigate the negative effects of O3 on plant physiology and growth based on a meta-analysis. We found that drought mitigated the negative effects of O3 on plant photosynthesis, but the modification of the O3 effect on the whole-plant biomass by drought was not significant. This is explained by a compensatory response of water-deficient plants that leads to increased metabolic costs. Relative to water control condition, reduced water treatment decreased the effects of O3 on photosynthetic traits, and leaf and root biomass in deciduous broadleaf species, while all traits in evergreen coniferous species showed no significant response. This suggested that the mitigating effects of drought on the negative impacts of O3 on the deciduous broadleaf species were more extensive than on the evergreen coniferous ones. Therefore, to avoid over- or underestimations when assessing the impact of O3 on vegetation growth, soil moisture should be considered. These results contribute to a better understanding of terrestrial ecosystem responses under global change.

Anthocyanins act as a sugar-buffer and an alternative electron sink in response to starch depletion during leaf senescence: a case study on a typical anthocyanic tree species, Acer japonicum.

We hypothesized that anthocyanins act as a sugar-buffer and an alternative electron sink during leaf senescence to prevent sugar-mediated early senescence and photoinhibition. To elucidate the role of anthocyanin, we monitored seasonal changes in photosynthetic traits, sugar, starch and N contents, pigment composition, and gene expression profiles in leaves exposed to substantially different light conditions within a canopy of an adult fullmoon maple (Acer japonicum) tree. Enhancement of starch amylolysis accompanied by cessation of starch synthesis occurred in the same manner independent of light conditions. Leaf sugar contents increased, but reached upper limits in the late stage of leaf senescence, even though leaf anthocyanins further increased after complete depletion of starch. Sun-exposed leaves maintained higher energy consumption via electron flow than shade-grown leaves during leaf N resorption. Thus, anthocyanins accumulated in sun-exposed leaves might have a regulative role as a sugar-buffer, retarding leaf senescence, and an indirect photoprotective role as an alternative sink for electron consumption to compensate declines in other metabolic processes such as starch and protein synthesis. In this context, anthocyanins may be key substrates protecting both outer-canopy leaves (against photoinhibition) and inner-canopy leaves (via shading by outer-canopy leaves) from high light stress during N resorption.

Ozone risk assessment with free-air controlled exposure (FACE) experiments: A critical revisit.

Since risk assessments of tropospheric ozone (O3) are crucial for agricultural and forestry sectors, there is a growing body for realistic assessments by a stomatal flux-based approach in Free-Air Controlled Exposure (FACE) facilities. Ozone risks are normally described as relative risks (RRs), which are calculated by assuming the biomass or yield at zero O3 dose as "reference". However, the estimation of the reference biomass or yield is challenging due to a lack of O3-clean-air treatment at the FACEs and the extrapolation without data in a low O3 range increases the bias for estimating the reference values. Here, we reviewed a current methodology for the risk assessment at FACEs and presented a simple and effective way ("modified Paoletti's approach") of defining RRs just using biomass or yield data with a range of expected impacts under the FACE conditions hypothesizing three possible scenarios based on prediction limits using 95% credible intervals (CI) (1. Best fit using the intercept as reference, 2. Optimistic scenario using a lower CI and 3. Worst scenario using an upper CI). As a result, O3-sensitive species show a relatively narrow effect range (optimistic vs. worst scenario) whereas a wide range of response may be possibly taken in resistant species. Showing a possible effect range allows for a comprehensive understanding of the potential risks and its uncertainties related to a species sensitivity to O3. As a supporting approach, we also recommend to use scientifically relevant tools (i.e., ethylenediurea treatments; mechanistic plant models) for strengthening the obtained results for the RRs against O3. Interestingly, the moderately sensitive or resistant species showed non-linear rather than linear dose-response relationships, suggesting a need for the flexible functional form for the risk assessment to properly describe the complex plant response such as hormesis, which depends on their plasticity to O3 stress.

Phenology- and light-based parameterization of stomatal conductance model for urban woody species in northern China.

Surface ozone (O3) poses a significant threat to urban vegetation health, and assessing the O3 risk across woody species is of vital importance for maintaining the health of urban infrastructure. In the present study, Jarvis-type stomatal conductance model was parameterized for ten urban species in northern China. Incorporating the effects of time of day and diurnal O3 concentration significantly enhanced the model performance. For different plant functional types (greening trees, greening shrubs, and orchard-grown trees), three parameterizations were established to estimate stomatal O3 uptake (POD1, phytotoxic O3 dose over an hourly threshold of 1 nmol m-2 s-1). The differences in POD1 between greening trees and shrubs were primarily due to the difference in their stomatal sensitivity to light. Orchard-grown trees displayed the lowest O3 removal capacity (lowest value of POD1) because of their shorter growing season despite of high stomatal conductance. These results indicated that plant phenology and light responsiveness determined stomatal O3 uptake, and the three parameterizations developed here could be applicable to various urban species in northern regions. Among climatic factors for O3 risk assessment, O3 concentration was the most important factor determining annual variation of POD1, which was primarily driven by air temperature. However, when O3 pollution decreased, O3 concentration exhibited less dependence on temperature and more dependence on light. These findings provide crucial insights for urban policy-makers and environmental scientists aiming to mitigate O3 pollution effects and enhance urban vegetation health.

The interplay of soil physicochemical properties, methanogenic diversity, and abundance governs methane production potential in paddy soil subjected to multi-decadal straw incorporation.

Straw incorporation holds significant promise for enhancing soil fertility and mitigating air pollution stemming from straw burning. However, this practice concurrently elevates the production and emission of methane (CH4) from paddy ecosystems. Despite its environmental impact, the precise mechanisms behind the heightened CH4 production resulting from long-term straw incorporation remain elusive. In a 32-year field experiment featuring three fertilization treatments (CFS-chemical fertilizer with wheat straw, CF-chemical fertilizer, and CK-unamended), we investigated the impact of abiotic (soil physicochemical properties) and biotic (methanogenic abundance, diversity, and community composition) factors on CH4 production in paddy fields. Results revealed a significantly higher CH4 production potential under CFS treatment compared to CF and CK treatments. The partial least squares path model revealed that soil physicochemical properties (path coefficient = 0.61), methanogenic diversity (path coefficient = -0.43), and methanogenic abundance (path coefficient = 0.29) collectively determined CH4 production potential, explaining 77% of the variance. Enhanced soil organic carbon content and water content, resulting from straw incorporation, emerged as pivotal factors positively correlated with CH4 production potential. Under CFS treatment, lower Shannon index of methanogens, compared to CF and CK treatments, was attributed to increased Methanosarcina. Notably, the Shannon index and relative abundance of Methanosarcina exhibited negative and positive correlations with CH4 production potential, respectively. Methanogenic abundance, bolstered by straw incorporation, significantly amplified overall potential. This comprehensive analysis underscores the joint influence of abiotic and biotic factors in regulating CH4 production potential during multi-decadal straw incorporation.

Flavonoids commonly induce hormetic responses.

The present paper provides a new perspective of previously published findings by Siwak (Food Chem 141:1227-1241, 2013) which showed that 15 structurally diverse flavonoids reduced toxicity (i.e., enhanced cell viability) from hypochlorite using the MTT assay within a pre-conditioning experimental protocol, with each agent showing a similar biphasic concentration response relationship. We use this Commentary to point out that each of the concentration response relationships are consistent with the hormetic dose response. The paper of Siwak (Food Chem 141:1227-1241, 2013) is unique in that it provides a comparison of a relatively large number of agents using the identical experimental protocol.

Soil Microbial Community Involved in Nitrogen Cycling in Rice Fields Treated with Antiozonant under Ambient Ozone.

Ethylenediurea (EDU) can effectively mitigate the crop yield loss caused by ozone (O3), a major, phytotoxic air pollutant. However, the relevant mechanisms are poorly understood, and the effect of EDU on soil ecosystems has not been comprehensively examined. In this study, a hybrid rice variety (Shenyou 63) was cultivated under ambient O3 and sprayed with 450 ppm EDU or water every 10 days. Real time quantitative polymerase chain reaction (RT-qPCR) showed that EDU had no significant effect on the microbial abundance in either rhizospheric or bulk soils. By applying both metagenomic sequencing and the direct assembly of nitrogen (N)-cycling genes, EDU was found to decrease the abundance of functional genes related to nitrification and denitrification processes. Moreover, EDU increased the abundance of genes involved in N-fixing. Although the abundance of some functional genes did not change significantly, nonmetric multidimensional scaling (NMDS) and a principal coordinates analysis (PCoA) suggested that the microbial community structure involved in N cycling was altered by EDU. The relative abundances of nifH-and norB-harboring microbial genera in the rhizosphere responded differently to EDU, suggesting the existence of functional redundancy, which may play a key role in sustaining microbially mediated N-cycling under ambient O3. IMPORTANCE Ethylenediurea (EDU) is hitherto the most efficient phytoprotectant agent against O3 stress. However, the underlying biological mechanisms of its mode of action are not clear, and the effects of EDU on the environment are still unknown, limiting its large-scale application in agriculture. Due to its sensitivity to environmental changes, the microbial community can be used as an indicator to assess the environmental impacts of agricultural practices on soil quality. This study aimed to unravel the effects of EDU spray on the abundance, community structure, and ecological functions of microbial communities in the rhizosphere of rice plants. Our study provides a deep insight into the impact of EDU spray on microbial-mediated N cycling and the structure of N-cycling microbial communities. Our findings help to elucidate the mode of action of EDU in alleviating O3 stress in crops from the perspective of regulating the structure and function of the rhizospheric soil microbial community.

Nitric oxide and hormesis.

This present paper provides an assessment of the occurrence of nitric oxide (NO)-induced hormetic-biphasic dose/concentration relationships in biomedical research. A substantial reporting of such NO-induced hormetic effects was identified with particular focus on wound healing, tumor promotion, and sperm biology, including mechanistic assessment and potential for translational applications. Numerous other NO-induced hormetic effects have been reported, but require more development prior to translational applications. The extensive documentation of NO-induced biphasic responses, across numerous organs (e.g., bone, cardiovascular, immune, intestine, and neuronal) and cell types, suggests that NO-induced biological activities are substantially mediated via hormetic processes. These observations are particularly important because broad areas of NO biology are constrained by the quantitative features of the hormetic response. This determines the amplitude and width of the low dose stimulation, affecting numerous biomedical implications, study design features (e.g., number of doses, dose spacing, sample sizes, statistical power), and the potential success of clinical trials.

Effect of ozone stress on crop productivity: A threat to food security.

Tropospheric ozone (O3), the most important phytotoxic air pollutant, can deteriorate crop quality and productivity. Notably, satellite and ground-level observations-based multimodel simulations demonstrate that the present and future predicted O3 exposures could threaten food security. Hence, the present study aims at reviewing the phytotoxicity caused by O3 pollution, which threatens the food security. The present review encompasses three major aspects; wherein the past and prevailing O3 concentrations in various regions were compiled at first, followed by discussing the physiological, biochemical and yield responses of economically important crop species, and considering the potential of O3 protectants to alleviate O3-induced phytotoxicity. Finally, the empirical data reported in the literature were quantitatively analysed to show that O3 causes detrimental effect on physiological traits, photosynthetic pigments, growth and yield attributes. The review on prevailing O3 concentrations over various regions, where economically important crop are grown, and their negative impact would support policy makers to implement air pollution regulations and the scientific community to develop countermeasures against O3 phytotoxicity for maintaining food security.

Sublethal chemical stimulation of arthropod parasitoids and parasites of agricultural and environmental importance.

An increasing number of studies have reported stimulation of various organisms in the presence of environmental contaminants. This has created a need to critically evaluate sublethal stimulation and hormetic responses of arthropod parasitoids and parasites following exposure to pesticides and other contaminants. Examining this phenomenon with a focus on arthropods of agricultural and environmental importance serves as the framework for this literature review. This review shows that several pesticides, with diverse chemical structures and different modes of action, applied individually or in combination at sublethal doses, commonly stimulate an array of arthropod parasitoids and parasites. Exposure at sublethal doses can enhance responses related to physiology (e.g., respiration, total lipid content, and total protein content), behavior (e.g., locomotor activity, antennal drumming frequency, host location, and parasitization), and fitness (longevity, growth, fecundity, population net and gross reproduction). Concordantly, the parasitic potential (e.g., infestation efficacy, parasitization rate, and parasitoid/parasite emergence) can be increased, and as a result host activities inhibited. There is some evidence illustrating hormetic dose-responses, but the relevant literature commonly included a limited number and range of doses, precluding a robust differentiation between sub- and superNOAEL (no-observed-adverse-effect level) stimulation. These results reveal a potentially significant threat to ecological health, through stimulation of harmful parasitic organisms by environmental contaminants, and highlight the need to include sublethal stimulation and hormetic responses in relevant ecological pesticide risk assessments. Curiously, considering a more utilitarian view, hormesis may also assist in optimizing mass rearing of biological control agents for field use, a possibility that also remains neglected.

Moringa induces its beneficial effect via hormesis.

Moringa oleifera, a traditional Indian herb, is widely known for its capacity to induce antioxidant, anti-inflammatory and other chemoprotective effects in a broad range of biomedical models. These perspectives have led to an extensive number of studies using various moringa extracts to evaluate its capacity to protect biological systems from oxidative stress and to explore whether it could be used to slow the onset of numerous age-related conditions and diseases. Moringa extracts have also been applied to prevent damage to plants from oxidative and saline stresses, following hormetic dose­response patterns. The present paper provides the first integrated and mechanistically based assessment showing that moringa extracts commonly induce hormetic dose responses and that many, perhaps most, of the beneficial effects of moringa are due to its capacity to act as an hormetic agent.

Hormesis: Transforming disciplines that rely on the dose response.

This article tells the story of hormesis from its conceptual and experimental origins, its dismissal by the scientific and medical communities in the first half of the 20th century, and its rediscovery over the past several decades to be a fundamental evolutionary adaptive strategy. The upregulation of hormetic adaptive mechanisms has the capacity to decelerate the onset and reduce the severity of a broad spectrum of common age-related health, behavioral, and performance decrements and debilitating diseases, thereby significantly enhancing the human health span. Incorporation of hormetic-based lifestyle options within the human population would have profoundly positive impacts on the public health, significantly reducing health care costs.

Elevated ozone inhibits isoprene emission of a diploid and a triploid genotype of Populus tomentosa by different mechanisms.

Ozone (O3) pollution affects plant growth and isoprene (ISO) emission. However, the response mechanism of isoprene emission rate (ISOrate) to elevated O3 (EO3) remains poorly understood. ISOrate was investigated in two genotypes (diploid and triploid) of Chinese white poplar (Populus tomentosa Carr.) exposed to EO3 in an open top chamber system. The triploid genotype had higher photosynthetic rate (A) and stomatal conductance (gs) than the diploid one. EO3 significantly decreased A, gs, and ISOrate of middle and lower leaves in both genotypes. In the diploid genotype, the reduction of ISOrate was caused by a systematic decrease related to ISO synthesis capacity, as indicated by decreased contents of the isoprene precursor dimethylallyl diphosphate and decreased isoprene synthase protein and activity. On the other hand, the negative effect of O3 on ISOrate of the triploid genotype did not result from inhibited ISO synthesis capacity, but from increased ISO oxidative loss within the leaf. Our findings will be useful for breeding poplar genotypes with high yield and lower ISOrate, depending on local atmospheric volatile organic compound/NOx ratio, to cope with both the rising O3 concentrations and increasing biomass demand. They can also inform the incorporation of O3 effects into process-based models of isoprene emission.

Strategic roadmap to assess forest vulnerability under air pollution and climate change.

Although it is an integral part of global change, most of the research addressing the effects of climate change on forests have overlooked the role of environmental pollution. Similarly, most studies investigating the effects of air pollutants on forests have generally neglected the impacts of climate change. We review the current knowledge on combined air pollution and climate change effects on global forest ecosystems and identify several key research priorities as a roadmap for the future. Specifically, we recommend (1) the establishment of much denser array of monitoring sites, particularly in the South Hemisphere; (2) further integration of ground and satellite monitoring; (3) generation of flux-based standards and critical levels taking into account the sensitivity of dominant forest tree species; (4) long-term monitoring of N, S, P cycles and base cations deposition together at global scale; (5) intensification of experimental studies, addressing the combined effects of different abiotic factors on forests by assuring a better representation of taxonomic and functional diversity across the ~73,000 tree species on Earth; (6) more experimental focus on phenomics and genomics; (7) improved knowledge on key processes regulating the dynamics of radionuclides in forest systems; and (8) development of models integrating air pollution and climate change data from long-term monitoring programs.

Hormesis: A General Biological Principle.

Hormesis drives biological modifications from cells to higher levels of biological organization and emerges as a general basic principle of biology, integrating evolution, ecology, medicine, physiology, toxicology, and public health.

Hormesis: Wound healing and keratinocytes.

Hormetic dose responses (i.e., a biphasic dose/concentration response characterized by a low dose stimulation and a high dose inhibition) are shown herein to be commonly reported in the dermal wound healing process, with the particular focus on cell viability, proliferation, and migration of human keratinocytes in in vitro studies. Hormetic responses are induced by a wide range of substances, including endogenous agents, numerous drug and nanoparticle preparations and especially plant derived extracts, including many well-known dietary supplements as well as physical stressor agents, such as low-level laser treatments. Detailed mechanistic studies have identified common signaling pathways and their cross-pathway communications that mediate the hormetic dose responses. These findings suggest that the concept of hormesis plays a fundamental role in wound healing, with important potential implications for agent screening and evaluation, as well as clinical strategies.

Hormesis: wound healing and fibroblasts.

Hormetic dose responses are reported here to occur commonly in the dermal wound healing process, with the particular focus on cell viability, proliferation, migration and collagen deposition of human and murine fibroblasts with in vitro studies. Hormetic responses were induced by a wide range of substances, including endogenous agents, pharmaceutical preparations, plant-derived extracts including many well-known dietary supplements, as well as physical stressor agents such as low-level laser treatments. Detailed mechanistic studies have identified common signaling pathways and their cross-pathway communications that mediate the hormetic dose responses. These findings complement and extend a similar comprehensive assessment concerning the occurrence of hormetic dose responses in keratinocytes. These findings demonstrate the generality of the hormetic dose response for key wound healing endpoints, suggesting that the hormesis concept has a fundamental role in wound healing, with respect to guiding strategies for experimental evaluation as well as therapeutic applications.

Hormesis is an evolutionary expectation: implications for aging.

This article argues that evolution and the concept of hormesis are biologically inseparable. It proposes that evolutionary processes led to the selection of inducible adaptive hormetic strategies that are necessary for wellbeing and survival. Hormesis has been demonstrated in essentially all organisms in which it has been studied from bacteria to humans, showing its highly conserved features. This evolution-hormesis integration should be a central feature in both understanding the biology of aging but also in ways to enhance improved health-based aging strategies.