Paired metabolomics and volatilomics provides insight into transient high light stress response mechanisms of the coral Montipora mollis.

The coral holobiont is underpinned by complex metabolic exchanges between different symbiotic partners, which are impacted by environmental stressors. The chemical diversity of the compounds produced by the holobiont is high and includes primary and secondary metabolites, as well as volatiles. However, metabolites and volatiles have only been characterised in isolation so far. Here, we applied a paired metabolomic-volatilomic approach to characterise holistically the chemical response of the holobiont under stress. Montipora mollis fragments were subjected to high-light stress (8-fold higher than the controls) for 30 min. Photosystem II (PSII) photochemical efficiency values were 7-fold higher in control versus treatment corals immediately following high-light exposure, but returned to pre-stress levels after 30 min of recovery. Under high-light stress, we identified an increase in carbohydrates (> 5-fold increase in arabinose and fructose) and saturated fatty acids (7-fold increase in myristic and oleic acid), together with a decrease in fatty acid derivatives in both metabolites and volatiles (e.g., 80% decrease in oleamide and nonanal), and other antioxidants (~ 85% decrease in sorbitol and galactitol). These changes suggest short-term light stress induces oxidative stress. Correlation analysis between volatiles and metabolites identified positive links between sorbitol, galactitol, six other metabolites and 11 volatiles, with four of these compounds previously identified as antioxidants. This suggests that these 19 compounds may be related and share similar functions. Taken together, our findings demonstrate how paired metabolomics-volatilomics may illuminate broader metabolic shifts occurring under stress and identify linkages between uncharacterised compounds to putatively determine their functions.

A new conceptual and quantitative approach to exploring and defining potential open-access olfactory information.

All organisms emit odour, providing 'open-access' olfactory information for any receiver with the right sensory apparatus. Characterizing open-access information emitted by groups of organisms, such as plant species, provides the means to answer significant questions about ecological interactions and their evolution. We present a new conceptual framework defining information reliability and a practical method to characterize and recover information from amongst olfactory noise. We quantified odour emissions from two tree species, one focal group and one outgroup, to demonstrate our approach using two new R statistical functions. We explore the consequences of relaxing or tightening criteria defining information and, from thousands of odour combinations, we identify and quantify those few likely to be informative. Our method uses core general principles characterizing information while incorporating knowledge of how receivers detect and discriminate odours. We can now map information in consistency-precision reliability space, explore the concept of information, and test information-noise boundaries, and between cues and signals.

Heat stress decreases the diversity, abundance and functional potential of coral gas emissions.

Terrestrial ecosystems emit large quantities of biogenic volatile organic compounds (BVOCs), many of which play important roles in abiotic stress responses, pathogen and grazing defences, inter- and intra-species communications, and climate regulation. Conversely, comparatively little is known about the diversity and functional potential of BVOCs produced in the marine environment, especially in highly productive coral reefs. Here we describe the first 'volatilomes' of two common reef-building corals, Acropora intermedia and Pocillopora damicornis, and how the functional potential of their gaseous emissions is altered by heat stress events that are driving rapid deterioration of coral reef ecosystems worldwide. A total of 87 BVOCs were detected from the two species and the chemical richness of both coral volatilomes-particularly the chemical classes of alkanes and carboxylic acids-decreased during heat stress by 41% and 62% in A. intermedia and P. damicornis, respectively. Across both coral species, the abundance of individual compounds changed significantly during heat stress, with the majority (>86%) significantly decreasing compared to control conditions. Additionally, almost 60% of the coral volatilome (or 52 BVOCs) could be assigned to four key functional groups based on their activities in other species or systems, including stress response, chemical signalling, climate regulation and antimicrobial activity. The total number of compounds assigned to these functions decreased significantly under heat stress for both A. intermedia (by 35%) and P. damicornis (by 64%), with most dramatic losses found for climatically active BVOCs in P. damicornis and antimicrobial BVOCs in A. intermedia. Together, our observations suggest that future heat stress events predicted for coral reefs will reduce the diversity, quantity and functional potential of BVOCs emitted by reef-building corals, potentially further compromising the healthy functioning of these ecosystems.

Range size and growth temperature influence Eucalyptus species responses to an experimental heatwave.

Understanding forest tree responses to climate warming and heatwaves is important for predicting changes in tree species diversity, forest C uptake, and vegetation-climate interactions. Yet, tree species differences in heatwave tolerance and their plasticity to growth temperature remain poorly understood. In this study, populations of four Eucalyptus species, two with large range sizes and two with comparatively small range sizes, were grown under two temperature treatments (cool and warm) before being exposed to an equivalent experimental heatwave. We tested whether the species with large and small range sizes differed in heatwave tolerance, and whether trees grown under warmer temperatures were more tolerant of heatwave conditions than trees grown under cooler temperatures. Visible heatwave damage was more common and severe in the species with small rather than large range sizes. In general, species that showed less tissue damage maintained higher stomatal conductance, lower leaf temperatures, larger increases in isoprene emissions, and less photosynthetic inhibition than species that showed more damage. Species exhibiting more severe visible damage had larger increases in heat shock proteins (HSPs) and respiratory thermotolerance (Tmax ). Thus, across species, increases in HSPs and Tmax were positively correlated, but inversely related to increases in isoprene emissions. Integration of leaf gas-exchange, isoprene emissions, proteomics, and respiratory thermotolerance measurements provided new insight into mechanisms underlying variability in tree species heatwave tolerance. Importantly, warm-grown seedlings were, surprisingly, more susceptible to heatwave damage than cool-grown seedlings, which could be associated with reduced enzyme concentrations in leaves. We conclude that species with restricted range sizes, along with trees growing under climate warming, may be more vulnerable to heatwaves of the future.

Visit, consume and quit: Patch quality affects the three stages of foraging.

Foraging is a three-stage process during which animals visit patches, consume food and quit. Foraging theory exploring relative patch quality has mostly focused on patch use and quitting decisions, ignoring the first crucial step for any forager: finding food. Yet, the decision to visit a patch is just as important as the decision to quit, as quitting theories can only be used if animals visit patches in the first place. Therefore, to better understand the foraging process and predict its outcomes, it is necessary to explore its three stages together. We used the common brushtail possum (Trichosurus vulpecula) as a model to investigate foraging decisions in response to food varying in quality. In particular, we tested whether patch nutritional quality affected the following: (1) patch visits; (2) behaviours at the patch during a foraging visit; and (3) patch quitting decisions (quantified using giving up density-GUD). Free-ranging possums were presented with diets varying in nitrogen content and concomitantly volatile organic compound (VOC) composition at feeding stations in the wild. We found that possums were able to distinguish between different quality foods from afar, despite the location of the diets changed daily. Possums used VOC (i.e. odour cues) emitted by the diets to find and select patches from a distance. High-quality diets with higher protein and lower fibre were visited more often and for longer. Possums spent more time foraging on diets high in nutritional content, resulting in lower GUDs. Our study provides important quantitative evidence that foraging efficiency plays out during all the three stages of the foraging process (i.e. visit, consume and quit), and demonstrates the significance of considering all these stages together in future studies and foraging models. Sensory cues such as food odours play a critical role in helping foragers, including mammalian herbivores, find high-quality food. This allows foragers to make quick, accurate and important decisions about food patches well before patch quitting decisions come into play.

Optimisation of fuel reduction burning regimes for carbon, water and vegetation outcomes.

Fire plays a critical role in biodiversity, carbon balance, soil erosion, and nutrient and hydrological cycles. While empirical evidence shows that fuel reduction burning can reduce the incidence, severity and extent of unplanned fires in Australia and elsewhere, the integration of environmental values into fire management operations is not well-defined and requires further research and development. In practice, the priority for fuel reduction burning is effective mitigation of risk to life and property. Environmental management objectives, including maintenance of high quality water, reduction of CO2 emissions and conservation of biodiversity can be constrained by this priority. We explore trade-offs between fuel reduction burning and environmental management objectives and propose a framework for optimising fuel reduction burning for environmental outcomes.

A model of plant isoprene emission based on available reducing power captures responses to atmospheric CO2.

We present a unifying model for isoprene emission by photosynthesizing leaves based on the hypothesis that isoprene biosynthesis depends on a balance between the supply of photosynthetic reducing power and the demands of carbon fixation. We compared the predictions from our model, as well as from two other widely used models, with measurements of isoprene emission from leaves of Populus nigra and hybrid aspen (Populus tremula × P. tremuloides) in response to changes in leaf internal CO2 concentration (C(i)) and photosynthetic photon flux density (PPFD) under diverse ambient CO2 concentrations (C(a)). Our model reproduces the observed changes in isoprene emissions with C(i) and PPFD, and also reproduces the tendency for the fraction of fixed carbon allocated to isoprene to increase with increasing PPFD. It also provides a simple mechanism for the previously unexplained decrease in the quantum efficiency of isoprene emission with increasing C(a). Experimental and modelled results support our hypothesis. Our model can reproduce the key features of the observations and has the potential to improve process-based modelling of isoprene emissions by land vegetation at the ecosystem and global scales.

Isoprene emission protects photosynthesis but reduces plant productivity during drought in transgenic tobacco (Nicotiana tabacum) plants.

Isoprene protects the photosynthetic apparatus of isoprene-emitting plants from oxidative stress. The role of isoprene in the response of plants to drought is less clear. Water was withheld from transgenic isoprene-emitting and non-emitting tobacco (Nicotiana tabacum) plants, to examine: the response of isoprene emission to plant water deficit; a possible relationship between concentrations of the drought-induced phytohormone abscisic acid (ABA) and isoprene; and whether isoprene affected foliar reactive oxygen species (ROS) and lipid peroxidation levels. Isoprene emission did not affect whole-plant water use, foliar ABA concentration or leaf water potential under water deficit. Compared with well-watered controls, droughted non-emitting plants significantly increased ROS content (31-46%) and lipid peroxidation (30-47%), concomitant with decreased operating and maximum efficiencies of photosystem II photochemistry and lower leaf and whole-plant water use efficiency (WUE). Droughted isoprene-emitting plants showed no increase in ROS content or lipid peroxidation relative to well-watered controls, despite isoprene emission decreasing before leaf wilting. Although isoprene emission protected the photosynthetic apparatus and enhanced leaf and whole-plant WUE, non-emitting plants had 8-24% more biomass under drought, implying that isoprene emission incurred a yield penalty.

Effect of Maternal Flavour Conditioning Combined with Organic and Inorganic Iron-Supplemented Creep Feed on Piglet Performance and Haemoglobin Status.

Iron injections are vital but imperfect against iron deficiency anaemia (IDA). This experiment explored the effects on piglets of maternal flavour conditioning and the voluntary intake of anise flavoured, iron-supplemented creep feed compared with iron injections. The experiment was a 2 × 2 factorial arrangement: ±maternal exposure to dietary anise flavour and ±intramuscular injections of piglets. Twenty-three sows and their litters (242 piglets) were randomly allocated to one of four treatments (n = 5 or 6 per treatment): no flavour plus no injection (NF + NI); no flavour plus iron injection (NF + I); flavour plus no injection (F + NI); and flavour plus iron injection (F + I). All piglets could access anise flavoured, iron-supplemented creep feed (organic and inorganic forms) from D2 of birth. Sow feed intake and milk anethole concentration, piglet body weight (BW) and average daily gain (ADG), creep feed disappearance, piglet behavioural time budgets, and piglet blood glucose and haemoglobin concentrations were determined. Over the four-week study, the only significant differences found were that iron-injected piglets had reduced blood glucose (p = 0.036) on D14 and that maternal flavour provision increased the frequency of piglet creep feed interaction (p = 0.023) and decreased the frequency of suckling events (p = 0.009). In summary, maternal flavour conditioning reduced piglet creep feed neophobia without influencing consumption. The supplementation of creep feed with iron and anise flavour to piglets under the conditions of this trial was effective in preventing IDA, regardless of exposure to maternal flavouring conditioning.

Olfactory misinformation provides refuge to palatable plants from mammalian browsing.

Mammalian herbivores browse palatable plants of ecological and economical value. Undesirable neighbours can reduce browsing to these plants by providing 'associational refuge', but they can also compete for resources. Here we recreated the informative odour emitted by undesirable plants. We then tested whether this odour could act as virtual neighbours, providing browsing refuge to palatable eucalyptus tree seedlings. We found that protection using this method was equivalent to protection provided by real plants. Palatable seedlings were 17-20 times more likely to be eaten by herbivores without virtual, or real, neighbours. Because many herbivores use plant odour to forage, virtual neighbours could provide a useful practical management approach to help protect valued plants.

Volatile isoprenoid emissions from plastid to planet.

Approximately 1-2% of net primary production by land plants is re-emitted to the atmosphere as isoprene and monoterpenes. These emissions play major roles in atmospheric chemistry and air pollution-climate interactions. Phenomenological models have been developed to predict their emission rates, but limited understanding of the function and regulation of these emissions has led to large uncertainties in model projections of air quality and greenhouse gas concentrations. We synthesize recent advances in diverse fields, from cell physiology to atmospheric remote sensing, and use this information to propose a simple conceptual model of volatile isoprenoid emission based on regulation of metabolism in the chloroplast. This may provide a robust foundation for scaling up emissions from the cellular to the global scale.

A unifying conceptual model for the environmental responses of isoprene emissions from plants.

BACKGROUND AND AIMS: Isoprene is the most important volatile organic compound emitted by land plants in terms of abundance and environmental effects. Controls on isoprene emission rates include light, temperature, water supply and CO2 concentration. A need to quantify these controls has long been recognized. There are already models that give realistic results, but they are complex, highly empirical and require separate responses to different drivers. This study sets out to find a simpler, unifying principle. METHODS: A simple model is presented based on the idea of balancing demands for reducing power (derived from photosynthetic electron transport) in primary metabolism versus the secondary pathway that leads to the synthesis of isoprene. This model's ability to account for key features in a variety of experimental data sets is assessed. KEY RESULTS: The model simultaneously predicts the fundamental responses observed in short-term experiments, namely: (1) the decoupling between carbon assimilation and isoprene emission; (2) a continued increase in isoprene emission with photosynthetically active radiation (PAR) at high PAR, after carbon assimilation has saturated; (3) a maximum of isoprene emission at low internal CO2 concentration (ci) and an asymptotic decline thereafter with increasing ci; (4) maintenance of high isoprene emissions when carbon assimilation is restricted by drought; and (5) a temperature optimum higher than that of photosynthesis, but lower than that of isoprene synthase activity. CONCLUSIONS: A simple model was used to test the hypothesis that reducing power available to the synthesis pathway for isoprene varies according to the extent to which the needs of carbon assimilation are satisfied. Despite its simplicity the model explains much in terms of the observed response of isoprene to external drivers as well as the observed decoupling between carbon assimilation and isoprene emission. The concept has the potential to improve global-scale modelling of vegetation isoprene emission.

Symbiont Identity Impacts the Microbiome and Volatilome of a Model Cnidarian-Dinoflagellate Symbiosis.

The symbiosis between cnidarians and dinoflagellates underpins the success of reef-building corals in otherwise nutrient-poor habitats. Alterations to symbiotic state can perturb metabolic homeostasis and thus alter the release of biogenic volatile organic compounds (BVOCs). While BVOCs can play important roles in metabolic regulation and signalling, how the symbiotic state affects BVOC output remains unexplored. We therefore characterised the suite of BVOCs that comprise the volatilome of the sea anemone Exaiptasia diaphana ('Aiptasia') when aposymbiotic and in symbiosis with either its native dinoflagellate symbiont Breviolum minutum or the non-native symbiont Durusdinium trenchii. In parallel, the bacterial community structure in these different symbiotic states was fully characterised to resolve the holobiont microbiome. Based on rRNA analyses, 147 unique amplicon sequence variants (ASVs) were observed across symbiotic states. Furthermore, the microbiomes were distinct across the different symbiotic states: bacteria in the family Vibrionaceae were the most abundant in aposymbiotic anemones; those in the family Crocinitomicaceae were the most abundant in anemones symbiotic with D. trenchii; and anemones symbiotic with B. minutum had the highest proportion of low-abundance ASVs. Across these different holobionts, 142 BVOCs were detected and classified into 17 groups based on their chemical structure, with BVOCs containing multiple functional groups being the most abundant. Isoprene was detected in higher abundance when anemones hosted their native symbiont, and dimethyl sulphide was detected in higher abundance in the volatilome of both Aiptasia-Symbiodiniaceae combinations relative to aposymbiotic anemones. The volatilomes of aposymbiotic anemones and anemones symbiotic with B. minutum were distinct, while the volatilome of anemones symbiotic with D. trenchii overlapped both of the others. Collectively, our results are consistent with previous reports that D. trenchii produces a metabolically sub-optimal symbiosis with Aiptasia, and add to our understanding of how symbiotic cnidarians, including corals, may respond to climate change should they acquire novel dinoflagellate partners.

Ecotoxicological effects of plastics on plants, soil fauna and microorganisms: A meta-analysis.

The interactions of plastics and soil organisms are complex and inconsistent observations on the effects of plastics on soil organisms have been made in published studies. In this study, we assessed the effects of plastic exposure on plants, fauna and microbial communities, with a meta-analysis. Using a total of 2936 observations from 140 publications, we analysed how responses in plants, soil fauna and microorganisms depended on the plastic concentration, size, type, species and exposure media. We found that overall plastics caused substantial detrimental effects to plants and fauna, but less so to microbial diversity and richness. Plastic concentration was one of the most important factors explaining variations in plant and faunal responses. Larger plastics (>1 µm) caused unfavourable changes to plant growth, germination and oxidative stress, while nanoplastics (NPs; ≤ 1 µm) only increased oxidative stress. On the contrary, there was a clear trend showing that small plastics adversely affected fauna reproduction, survival and locomotion than large plastics. Plant responses were indifferent to plastic type, with most studies conducted using polyethylene (PE) and polystyrene (PS) plastics, but soil fauna were frequently more sensitive to PS than to PE exposure. Plant species played a vital role in some parameters, with the effects of plastics being considerably greater on vegetable plants than on cereal plants.

Isoprene synthesis in plants: lessons from a transgenic tobacco model.

Isoprene is a highly reactive gas, and is emitted in such large quantities from the biosphere that it substantially affects the oxidizing potential of the atmosphere. Relatively little is known about the control of isoprene emission at the molecular level. Using transgenic tobacco lines harbouring a poplar isoprene synthase gene, we examined control of isoprene emission. Isoprene synthase required chloroplastic localization for catalytic activity, and isoprene was produced via the methyl erythritol (MEP) pathway from recently assimilated carbon. Emission patterns in transgenic tobacco plants were remarkably similar to naturally emitting plants under a wide variety of conditions. Emissions correlated with photosynthetic rates in developing and mature leaves, and with the amount of isoprene synthase protein in mature leaves. Isoprene synthase protein levels did not change under short-term increase in heat/light, despite an increase in emissions under these conditions. A robust circadian pattern could be observed in emissions from long-day plants. The data support the idea that substrate supply and changes in enzyme kinetics (rather than changes in isoprene synthase levels or post-translational regulation of activity) are the primary controls on isoprene emission in mature transgenic tobacco leaves.

Phloem sap metabolites vary according to the interactive effects of nutrient supply and seasonal conditions in Eucalyptus globulus (Labill).

Improving the efficiency of fertilizer application is paramount to both the sustainability and profitability of forest plantations. Therefore, developing reliable, cost-effective tools to assess tree nutritional status is of great interest. This investigation sought to assess the use of phloem sap-derived metabolites as an indicator of nutritional status on a background of seasonal water availability of Eucalyptus globulus (Labill) trees grown under field conditions. Phloem is a central conduit for long-distance transport and signaling in plants and offers great promise in reflecting plant-scale resource limitations. Changes in the abundance of solutes and isotopes in phloem sap are sensitive to environmental cues. With a focus on both water and nutrient availability, we characterize patterns in phloem sugars, amino acids and the abundance of carbon isotopes in phloem sap obtained from E. globulus among different seasons and fertilizer treatments. Phloem-derived total amino acid concentration was found to increase with an increasing nitrogen (N) supply; however, this response was lost with the concurrent addition of phosphorus and at the highest level of N supply. Significant seasonal variation in all measured parameters was also detected, highlighting the need for caution in making quantitative relationships with growth. Broader implications of the interactive effects of both water supply and multi-nutrient additions and relationships with growth are discussed.

Genetic structure and regulation of isoprene synthase in Poplar (Populus spp.).

Isoprene is a volatile 5-carbon hydrocarbon derived from the chloroplastic methylerythritol 2-C-methyl-D: -erythritol 4-phosphate isoprenoid pathway. In plants, isoprene emission is controlled by the enzyme isoprene synthase; however, there is still relatively little known about the genetics and regulation of this enzyme. Isoprene synthase gene structure was analysed in three poplar species. It was found that genes encoding stromal isoprene synthase exist as a small gene family, the members of which encode virtually identical proteins and are differentially regulated. Accumulation of isoprene synthase protein is developmentally regulated, but does not differ between sun and shade leaves and does not increase when heat stress is applied. Our data suggest that, in mature leaves, isoprene emission rates are primarily determined by substrate (dimethylallyl diphosphate, DMADP) availability. In immature leaves, where isoprene synthase levels are variable, emission levels are also influenced by the amount of isoprene synthase protein. No thylakoid isoforms could be identified in Populus alba or in Salix babylonica. Together, these data show that control of isoprene emission at the genetic level is far more complicated than previously assumed.

Effects of fosmidomycin on plant photosynthesis as measured by gas exchange and chlorophyll fluorescence.

In higher plants, many isoprenoids are synthesised via the chloroplastic 1-deoxy-D-xylulose 5-phosphate/2-C-methyl-D-erythritol 4-phosphate (MEP) pathway. Attempts to elucidate the function of individual isoprenoids have used the antibiotic/herbicidal compound fosmidomycin (3-[N-formyl-N-hydroxy amino] propyl phosphonic acid) to inhibit this pathway. Examination of the effect of fosmidomycin on the major components of photosynthesis in leaves of white poplar (Populus alba) and tobacco (Nicotiana tabacum) was made. Fosmidomycin reduced net photosynthesis in both species within 1 h of application, but only when photosynthesis was light-saturated. In P. alba, these reductions were confounded by high light and fosmidomycin inducing stomatal patchiness. In tobacco, this was caused by significant reductions in PSII chlorophyll fluorescence and reductions in V(cmax) and J(max). Our data indicate that the diminution of photosynthesis is likely a complex effect resulting from the inhibition of multiple MEP pathway products, resulting in photoinhibition and photo-damage. These effects should be accounted for in experimental design and analysis when using fosmidomycin to avoid misinterpretation of results as measured by gas exchange and chlorophyll fluorescence.

Gas exchange and photosynthetic performance of the tropical tree Acacia nigrescens when grown in different CO(2) concentrations.

The photosynthetic responses of the tropical tree species Acacia nigrescens Oliv. grown at different atmospheric CO(2) concentrations-from sub-ambient to super-ambient-have been studied. Light-saturated rates of net photosynthesis (A (sat)) in A. nigrescens, measured after 120 days exposure, increased significantly from sub-ambient (196 microL L(-1)) to current ambient (386 microL L(-1)) CO(2) growth conditions but did not increase any further as [CO(2)] became super-ambient (597 microL L(-1)). Examination of photosynthetic CO(2) response curves, leaf nitrogen content, and leaf thickness showed that this acclimation was most likely caused by reduction in Rubisco activity and a shift towards ribulose-1,5-bisphosphate regeneration-limited photosynthesis, but not a consequence of changes in mesophyll conductance. Also, measurements of the maximum efficiency of PSII and the carotenoid to chlorophyll ratio of leaves indicated that it was unlikely that the pattern of A (sat) seen was a consequence of growth [CO(2)] induced stress. Many of the photosynthetic responses examined were not linear with respect to the concentration of CO(2) but could be explained by current models of photosynthesis.

Isoprene synthesis protects transgenic tobacco plants from oxidative stress.

Isoprene emission represents a significant loss of carbon to those plant species that synthesize this highly volatile and reactive compound. As a tool for studying the role of isoprene in plant physiology and biochemistry, we developed transgenic tobacco plants capable of emitting isoprene in a similar manner to and at rates comparable to a naturally emitting species. Thermotolerance of photosynthesis against transient high-temperature episodes could only be observed in lines emitting high levels of isoprene; the effect was very mild and could only be identified over repetitive stress events. However, isoprene-emitting plants were highly resistant to ozone-induced oxidative damage compared with their non-emitting azygous controls. In ozone-treated plants, accumulation of toxic reactive oxygen species (ROS) was inhibited, and antioxidant levels were higher. Isoprene-emitting plants showed remarkably decreased foliar damage and higher rates of photosynthesis compared to non-emitting plants immediately following oxidative stress events. An inhibition of hydrogen peroxide accumulation in isoprene-emitting plants may stall the programmed cell death response which would otherwise lead to foliar necrosis. These results demonstrate that endogenously produced isoprene provides protection from oxidative damage.