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Over the past decades, introduction of many fast-growing hygrophilic, and economically valuable plants into xeric environments has occurred. However, production and even survival of these species may be threatened by harsh climatic conditions unless an effective physiological and metabolic plasticity is available. Moringa oleifera Lam. a multipurpose tree originating from humid sub-tropical regions of India, is widely cultivated in many arid countries because of its multiple uses. We tested whether M. oleifera can adjust primary and secondary metabolism to efficiently cope with increasing water stress. It is shown that M. oleifera possesses an effective isohydric behavior. Water stress induced a quick and strong stomatal closure, driven by abscisic acid (ABA) accumulation, and leading to photosynthesis inhibition with consequent negative effects on biomass production. However, photochemistry was not impaired and maximal fluorescence and saturating photosynthesis remained unaffected in stressed leaves. We report for the first time that M. oleifera produces isoprene, and show that isoprene emission increased three-fold during stress progression. It is proposed that higher isoprene biosynthesis helps leaves cope with water stress through its antioxidant or membrane stabilizing action, and also indicates a general MEP (methylerythritol 4-phosphate) pathway activation that further helps protect photosynthesis under water stress. Increased concentrations of antioxidant flavonoids were also observed in water stressed leaves, and probably cooperate in limiting irreversible effects of the stress in M. oleifera leaves. The observed metabolic and phenotypic plasticity may facilitate the establishment of M. oleifera in xeric environments, sustaining the economic and environmental value of this plant. [ABSTRACT FROM AUTHOR]

Main conclusion AM symbiosis did not strongly affect Arundo donax performances under salt stress, although differences in the plants inoculated with two different fungi were recorded. The mechanisms at the basis of the improved tolerance to abiotic stresses by arbuscular mycorrhizal (AM) fungi have been investigated mainly focusing on food crops. In this work, the potential impact of AM symbiosis on the performance of a bioenergy crop, Arundo donax, under saline conditions was considered. Specifically, we tried to understand whether AM symbiosis helps this fast-growing plant, often widespread in marginal soils, withstand salt. A combined approach, involving eco-physiological, morphometric and biochemical measurements, was used and the effects of two different AM fungal species (Funneliformis mosseae and Rhizophagus irregularis) were compared. Results indicate that potted A. donax plants do not suffer permanent damage induced by salt stress, but photosynthesis and growth are considerably reduced. Since A. donax is a high-yield biomass crop, reduction of biomass might be a serious agronomical problem in saline conditions. At least under the presently experienced growth conditions, and plant-AM combinations, the negative effect of salt on plant performance was not rescued by AM fungal colonization. However, some changes in plant metabolisms were observed following AM-inoculation, including a significant increase in proline accumulation and a trend toward higher isoprene emission and higher H2O2, especially in plants colonized by R. irregularis. This suggests that AM fungal symbiosis influences plant metabolism, and plant-AM fungus combination is an important factor for improving plant performance and productivity, in presence or absence of stress conditions. [ABSTRACT FROM AUTHOR]

Isoprene is synthesized through the 2-C-methylerythritol-5-phosphate (MEP) pathway that also produces abscisic acid (ABA). Increases in foliar free ABA concentration during drought induce stomatal closure and may also alter ethylene biosynthesis. We hypothesized a role of isoprene biosynthesis in protecting plants challenged by increasing water deficit, by influencing ABA production and ethylene evolution. We performed a split-root experiment on Populus nigra L. subjected to three water treatments: well-watered (WW) plants with both root sectors kept at pot capacity, plants with both root compartments allowed to dry for 5 days (DD) and plants with one-half of the roots irrigated to pot capacity, while the other half did not receive water (WD). WD and WW plants were similar in photosynthesis, water relations, foliar ABA concentration and isoprene emission, whereas these parameters were significantly affected in DD plants: leaf isoprene emission increased despite the fact that photosynthesis declined by 85% and the ABA-glucoside/free ABA ratio decreased significantly. Enhanced isoprene biosynthesis in water-stressed poplars may have contributed to sustaining leaf ABA biosynthesis by keeping the MEP pathway active. However, this enhancement in ABA was accompanied by no change in ethylene biosynthesis, likely confirming the antagonistic role between ABA and ethylene. These results may indicate a potential cross-talk among isoprene, ABA and ethylene under drought. [ABSTRACT FROM AUTHOR]

The functional reasons for isoprene emission are still a matter of hot debate. It was hypothesized that isoprene biosynthesis evolved as an ancestral mechanism in plants adapted to high water availability, to cope with transient and recurrent oxidative stresses during their water-to-land transition. There is a tight association between isoprene emission and species hygrophily, suggesting that isoprene emission may be a favorable trait to cope with occasional exposure to stresses in mesic environments. The suite of morpho-anatomical traits does not allow a conservative water use in hygrophilic mesophytes challenged by the environmental pressures imposed or exacerbated by drought and heat stress. There is evidence that in stressed plants the biosynthesis of isoprene is uncoupled from photosynthesis. Because the biosynthesis of isoprene is costly, the great investment of carbon and energy into isoprene must have relevant functional reasons. Isoprene is effective in preserving the integrity of thylakoid membranes, not only through direct interaction with their lipid acyl chains, but also by up-regulating proteins associated with photosynthetic complexes and enhancing the biosynthesis of relevant membrane components, such as mono- and di-galactosyl-diacyl glycerols and unsaturated fatty acids. Isoprene may additionally protect photosynthetic membranes by scavenging reactive oxygen species. Here we explore the mode of actions and the potential significance of isoprene in the response of hygrophilic plants when challenged by severe stress conditions associated to rapid climate change in temperate climates, with special emphasis to the concomitant effect of drought and heat. We suggest that isoprene emission may be not a good estimate for its biosynthesis and concentration in severely droughted leaves, being the internal concentration of isoprene the important trait for stress protection. [ABSTRACT FROM AUTHOR]

Genetic analyses have suggested that the clonal reproduction of Arundo donax has resulted in low genetic diversity. However, an earlier common garden phenotyping experiment identified specimens of A. donax with contrasting biomass yields (ecotypes 6 and 20). We utilized the same well-established stands to investigate the photosynthetic and stress physiology of the A. donax ecotypes under irrigated and drought conditions. Ecotype 6 produced the largest yields in both treatments. The A. donax ecotypes exhibited identical high leaf-level rates of photosynthesis ( PN) and stomatal conductance ( Gs) in the well-watered treatment. Soil drying induced reductions in PN and Gs, decreased use of light energy for photochemistry, impaired function of photosystem II and increased heat dissipation similarly in the two ecotypes. Levels of biologically active free-abscisic acid ( ABA) and fixed glycosylated- ABA increased earlier in response to the onset of water deficit in ecotype 6; however, as drought progressed, the ecotypes showed similar increases in both forms of ABA. This may suggest that because of the low genetic variability in A. donax the genes responding to drought might have been activated similarly in the two ecotypes, resulting in identical physiological responses to water deficit. Despite the lack of physiological ecotypic differences that could be associated with yield, A. donax retained a high degree of PN and biomass gain under water deficit stress conditions. This may enable utilization of A. donax as a fast growing biomass crop in rain-fed marginal lands in hot drought prone climates. [ABSTRACT FROM AUTHOR]

Isoprene emission by terrestrial plants is believed to play a role in mitigating the effects of abiotic stress on photosynthesis. Ultraviolet-B light (UV-B) induces damage to the photosynthetic apparatus of plants, but the role of isoprene in UV-B tolerance is poorly understood. To investigate this putative protective role, we exposed non-emitting (NE) control and transgenic isoprene emitting (IE) Nicotiana tabacum (tobacco) plants to high intensity UV-B exposure. Methanol emissions increased with UV-B intensity, indicating oxidative damage. However, isoprene emission was unaffected during exposure to UV-B radiation, but declined in the 48 h following UV-B treatment at the highest UV-B intensities of 9 and 15 W m-2. Photosynthesis and the performance of photosystem II (PSII) declined to similar extents in IE and NE plants following UV-B exposure, suggesting that isoprene emission did not ameliorate the immediate impact of UV-B on photosynthesis. However, after the stress, photosynthesis and PSII recovered in IE plants, which maintained isoprene formation, but not in NE plants. Recovery of IE plants was also associated with elevated antioxidant levels and cycling; suggesting that both isoprene formation and antioxidant systems contributed to reinstating the integrity and functionality of cellular membranes and photosynthesis following exposure to excessive levels of UV-B radiation. [ABSTRACT FROM AUTHOR]

Water availability is a major limiting factor on plant growth and productivity. Considering that Eucalyptus spp. are among the few plant species able to produce both isoprene and monoterpenes, experiments were designed to investigate the response of isoprene emission and isoprenoid concentrations in Eucalyptus citriodora saplings exposed to decreasing fraction of transpirable soil water (FTSW). In particular, this study aimed to assess: (a) the kinetic of water stress-induced variations in photosynthesis, isoprene emission, and leaf isoprenoid concentrations during progressive soil water shortage as a function of FTSW; (b) the ultradian control of isoprene emission and photosynthesis under limited soil water availability; and (c) the optimum temperature sensitivity of isoprene emission and photosynthesis under severe water stress. The optimum temperature for isoprene emission did not change under progressive soil water deficit. However, water stress induced a reallocation of carbon through the MEP/DOXP pathway resulting in a qualitative change of the stored isoprenoids. The ultradian trend of isoprene emission was also unaffected under water stress, and a similar ultradian trend of stomatal and mesophyll conductances was also observed, highlighting a tight coordination between diffusion limitations to photosynthesis during water stress. The kinetics of photosynthetic parameters and isoprene emission in response to decreasing FTSW in E. citriodora are strikingly similar to those measured in other plant functional types. These findings may be useful to refine the algorithms employed in process-based models aiming to precisely up-scale carbon assimilation and isoprenoid emissions at regional and global scales. [ABSTRACT FROM AUTHOR]

The effects of the interaction between high growth temperatures and water stress on gas-exchange properties of Populus nigra saplings were investigated. Water stress was expressed as a function of soil water content (SWC) or fraction of transpirable soil water (FTSW). Isoprene emission and photosynthesis (A) did not acclimate in response to elevated temperature, whereas dark (Rn) and light (Rd) respiration underwent thermal acclimation. Rd was ∼30% lower than Rn irrespective of growth temperature and water stress level. Water stress induced a sharp decline, but not a complete inhibition, of both Rn and Rd. There was no significant effect of high growth temperature on the responses of A, stomatal conductance (gs), isoprene emission, Rn or Rd to FTSW. High growth temperature resulted in a significant increase in the SWC endpoint. Photosynthesis was limited mainly by CO2 acquisition in water-stressed plants. Impaired carbon metabolism became apparent only at the FTSW endpoint. Photosynthesis was restored in about a week following rewatering, indicating transient biochemical limitations. The kinetics of isoprene emission in response to FTSW confirmed that water stress uncouples the emission of isoprene from A, isoprene emission being unaffected by decreasing gs. The different kinetics of A, respiration and isoprene emission in response to the interaction between high temperature and water stress led to rising Rd/A ratio and amount of carbon lost as isoprene. Since respiration and isoprene sensitivity are much lower than A sensitivity to water stress, temperature interactions with water stress may dominate poplar acclimatory capability and maintenance of carbon homeostasis under climate change scenarios. Furthermore, predicted temperature increases in arid environments may reduce the amount of soil water that can be extracted before plant gas exchange decreases, exacerbating the effects of water stress even if soil water availability is not directly affected. [ABSTRACT FROM AUTHOR]

Black poplar ( Populus nigra L.) plants grown at 25 and 35°C were subjected to drought stress to assess the combined impact of two consequences of global climate change – rising temperature and drought – on isoprene biosynthesis and emission. At both temperatures, photosynthesis was inhibited by moderate drought, but isoprene emission only decreased when drought was prolonged. The mRNA transcript level, protein concentration and activity of isoprene synthase (ISPS) changed in concert with isoprene emission during drought stress. However, ISPS activity decreased before isoprene emission during drought development, indicating a tighter control of the emission at a transcriptional or post-transcriptional level under moderate drought stress, and at both temperatures. A residual isoprene emission was measured when photosynthesis was totally inhibited after 35 days of drought. This photosynthesis-independent emission of isoprene was probably dependent on a cytosolic carbon supply as all the properties of ISPS were drastically inhibited. Isoprene emission was associated with dark respiration during the entire drought stress period, and at both temperatures, indicating that the two processes are sustained by, but do not compete for, the same carbon source. Isoprene emission was directly related to phosphoenolpyruvate carboxylase activity in plants grown at 25°C and inversely related in plants grown at 35°C, suggesting a strong temperature control on the regulation of the pyruvate flowing from the cytosol to the plastidic isoprenoid biosynthetic pathway under drought stress and recovery. In re-watered plants, the temperature control on isoprene emission was suppressed, despite complete recovery of photosynthesis and ISPS activity similar to levels in plants subjected to mild drought stress. Our results reveal the overriding effects of drought on isoprene emission, possibly affecting protein level or substrate supply. These effects may largely offset the predicted impact of rising temperatures on the emission of isoprene in terrestrial ecosystems. [ABSTRACT FROM AUTHOR]

• The mechanism uncoupling isoprene emission and photosynthesis under drought was investigated in Populus alba saplings. • Isoprene emission, incorporation of 13C into the isoprene molecule, isoprene synthase (ISPS) activity, concentration and gene expression, and photosynthesis were measured as a function of the fraction of transpirable soil water (FTSW) and in plants recovering from drought. • Photosynthesis sharply declined below FTSW30 (a FTSW of 30%) and its inhibition was not caused by metabolic factors. A decline in isoprene emission was only evident towards the FTSW endpoint. 13C incorporation into isoprene was lower when photosynthesis was constrained by drought. ISPS activity was inhibited by mild drought, while ISPS gene expression and concentration declined in concert with isoprene emission at the FTSW endpoint. Following rewatering, isoprene emission was higher than, and photosynthesis was similar to, prestress values. ISPS activity and concentration, and 13C incorporation into isoprene, also rapidly recovered to prestress values, while ISPS gene expression remained low in rewatered plants. • Our experiment revealed a larger contribution of alternative carbon sources to isoprene emission only when photosynthesis was dramatically constrained by drought. Isoprene emission was likely controlled at the posttranscriptional level under severe drought. [ABSTRACT FROM AUTHOR]

In past studies, it was hypothesized that reductions in chloroplast isoprene emissions at high atmospheric CO2 concentrations were caused by competition between cytosolic and mitochondrial processes for the same substrate, possibly phosphoenolpyruvate (PEP). We conducted field and laboratory experiments using leaves of white poplar ( Populus alba L.) to identify whether an inverse relationship occurs between the dark respiration rate (a mitochondrial process) and the isoprene emission rate. Field experiments that were carried out in a free-air CO2-enriched (FACE) facility showed no clear effect of elevated CO2 on either isoprene emission rate or respiration rate by leaves. In young, not yet fully expanded leaves, low isoprene emission and high dark respiration rates were measured in both ambient and elevated CO2. In these leaves, isoprene emission was inversely correlated with dark respiration. It is possible to interpret from these results that, in young leaves, high rates of growth respiration compete with isoprene biosynthesis for the same substrate. However, it is also possible that the negative correlation reflects the contrasting reductions in growth respiration and increases in expression of the enzyme isoprene synthase at this final stage of leaf maturation. In contrast to our observations on young leaves, respiration rate and isoprene emission rate were positively correlated in older, fully expanded leaves (8 and 11 from apex). A positive correlation was also found between respiration rate and isoprene emission rate when these parameters were modulated using different ozone exposure, growth light intensity, growth temperature and exposure to different leaf temperatures in laboratory experiments. These data show that competition for substrate between isoprene biosynthesis and leaf respiration does not determine the rate of isoprene emission in most circumstances that affect both processes. A negative correlation was observed across all experiments between isoprene emission rate and the activity of phosphoenolpyruvate carboxylase (PEPc), a cytosolic enzyme that competes with isoprene biosynthesis for substrate. The cytosolic metabolite, PEP, occurs at a metabolic branch point from which substrate flows into three processes: (1) the production of pyruvate for mitochondrial respiration, (2) the production of oxaloacetate (OAA) by PEPc for anabolic support of mitochondrial respiration and (3) transport into the chloroplast to support chloroplastic demands for pyruvate, including isoprenoid biosynthesis. The results of our observations suggest that only the second process competes for substrate with isoprenoid synthesis, while the partitioning of PEP between mitochondrial respiration and chloroplast isoprenoid biosynthesis is controlled in a way that retains balance in substrate demand. [ABSTRACT FROM AUTHOR]

The direct and indirect impact of ozone on Populus alba was studied by exposing leaves enclosed in specially designed cuvettes for 30 days to high ozone (150 ppb, 11 h per day), while leaves developing above the cuvettes were exposed to ambient ozone. Gas exchanges and histo-anatomical parameters were measured to specifically understand whether ozone indirectly affects the anatomy and physiology of leaves. Three leaf classes were investigated: (1) those expanding above the cuvettes (A leaves); (2) those already developed inside the cuvettes (B leaves) and (3) those developing inside the cuvettes, since the beginning of the ozone treatment (C leaves). The anatomy and morphology of the first leaf developing outside the cuvette (A1) were strongly affected by ozone, whereas photosynthesis was not perturbed. However, in leaves of ozone-treated plants developing after A1, a large reduction of starch accumulation was observed, which suggests a delayed biosynthesis, or a very rapid export of starch toward other sinks. Isoprene emission was higher and isoprene synthase messenger RNA was more expressed in ozone-treated A1 leaves than in control leaves with similar ontogeny. This indicates that isoprene synthesis is stimulated by ozone, and reveals that isoprene emission is controlled at a transcriptional level. Leaves already developed inside the cuvette (B leaves) rapidly sensed ozone stress, which inhibited photosynthesis, stomatal conductance and isoprene emission. The observation that new leaves were developing inside the cuvettes during the treatment (C leaves) suggests that resistance to ozone may be acquired by plants. Leaves C showed a more packed and thinner mesophyll than controls of similar development, which may help reduce ozone penetration inside cells. They also showed a lower photosynthesis in comparison to controls and to other leaf classes, probably because of ribulose 1,5-bisphosphate carboxylase/oxygenase activity limitation, as inferred from photosynthesis response to intercellular CO2. However, isoprene emission was slightly stimulated also in C leaves, confirming that a large fraction of carbon is invested into isoprene formation under ozone stress. [ABSTRACT FROM AUTHOR]

• Co-inoculation with two PGPR strains has a potential for biotechnological application in areas affected by the drought. • PGPR strains with ACC deaminase activity improve total biomass and WUE under water deficit. • Inoculated plants show higher carbon assimilation during water stress progression. • Inoculation promotes isoprene emission, enhancing plant tolerance to water stress. Agricultural sustainability is an increasing need considering the challenges posed by climate change and rapid human population growth. The use of plant growth-promoting rhizobacteria (PGPR) may represent an excellent, new agriculture practice to improve soil quality while promoting growth and yield of important crop species subjected to water stress conditions. In this study, two PGPR strains with 1-Aminocyclopropane-1-Carboxylate (ACC) deaminase activity were co-inoculated in velvet bean plants to verify the physiological, biochemical and molecular responses to progressive water stress. The results of our study show that the total biomass and the water use efficiency of inoculated plants were higher than uninoculated plants at the end of the water stress period. These positive effects may be derived from a lower root ACC content (−45 %) in water-stressed inoculated plants than in uninoculated ones resulting in lower root ethylene emission. Furthermore, the ability of inoculated plants to maintain higher levels of both isoprene emission, a priming compound that may help to protect leaves from oxidative damage, and carbon assimilation during water stress progression may indicate the underlining metabolic processes conferring water stress tolerance. Overall, the experimental results show that co-inoculation with ACC deaminase PGPR positively affects tolerance to water deficit, confirming the potential for biotechnological applications in water-stressed agricultural areas. [ABSTRACT FROM AUTHOR]

Abstract: Acclimatory and direct responses of photosynthesis (A), stomatal (g s) and mesophyll (gm ) conductance, light (R L) and dark (R D) respiration, and isoprene emission, measured at different temperatures, were compared in one-year-old Populus × euramericana saplings grown at 25°C, 35°C, and in the saplings grown initially at 35°C and then exposed for two weeks at 25°C. Results show that A, g s and gm were significantly lower at 35°C, than at the other growth temperatures. This resulted in a downward acclimation of these parameters over the range of measuring temperatures. Both R L and R D also showed a considerable downward acclimation. However, the respiration to photosynthesis ratio increased with high temperatures, for R L and R D were more responsive to increasing growth temperatures than A at all measuring temperatures. This type of acclimation would lead to a shift in the carbon balance between photosynthesis and respiration under changing climatic conditions. Isoprene emission was greatly affected by temperature treatments and showed a downward acclimation to higher temperatures. Respiration and isoprene emission rates were directly related, independently of growth and measuring temperatures. These findings may be likely relevant to predict the emissions of isoprenoid in globally changing environmental conditions. [ABSTRACT FROM AUTHOR]