Winter survival in red clover: experimental evidence for interactions among stresses.

BACKGROUND: There is a lack of knowledge on the combined effects of different stresses on plants, in particular different stresses that occur during winter in temperate climates. Perennial herbaceous plants in temperate regions are exposed to many different stresses during winter, but except for the fact that cold temperatures induce resistance to a number of them, very little is known about their interaction effects. Knowledge about stress interactions is needed in order to predict effects of climate change on both agricultural production and natural ecosystems, and to develop adaptation strategies, e.g., through plant breeding. Here, we conducted a series of experiments under controlled conditions to study the interactions between cold (low positive temperature), clover rot infection (caused by Sclerotinia trifoliorum) and freezing, in red clover (Trifolium pratense) accessions. We also compared our results with winter survival in field experiments and studied associations between stress and shoot growth. RESULTS: Exposure to low positive temperatures (cold acclimation) induced resistance to clover rot. There was a clear negative interaction effect between freezing stress and clover rot infection, resulting in up to 37% lower survival rate compared to what would have been expected from the additive effect of freezing and infection alone. Freezing tolerance could continue to improve during incubation under artificial snow cover at 3 °C in spite of darkness, and we observed compensatory shoot growth following freezing after prolonged incubation. At the accession level, resistance to clover rot was negatively correlated with growth in the field during the previous year at a Norwegian location. It was also negatively correlated with the shoot regrowth of control plants after incubation. Clover rot resistance tests under controlled conditions showed limited correlation with clover rot resistance observed in the field, suggesting that they may reveal variation in more specific resistance mechanisms. CONCLUSIONS: We here demonstrate, for the first time, a strong negative interaction between freezing and infection with a winter pathogen. We also characterize the effects of cold acclimation and incubation in darkness at different temperatures on winter stress tolerance, and present data that support the notion that annual cycles of growth and stress resistance are associated at the genetic level.

Long-term drought effects on the thermal sensitivity of Amazon forest trees.

The continued functioning of tropical forests under climate change depends on their resilience to drought and heat. However, there is little understanding of how tropical forests will respond to combinations of these stresses, and no field studies to date have explicitly evaluated whether sustained drought alters sensitivity to temperature. We measured the temperature response of net photosynthesis, foliar respiration and the maximum quantum efficiency of photosystem II (Fv /Fm ) of eight hyper-dominant Amazonian tree species at the world's longest-running tropical forest drought experiment, to investigate the effect of drought on forest thermal sensitivity. Despite a 0.6°C-2°C increase in canopy air temperatures following long-term drought, no change in overall thermal sensitivity of net photosynthesis or respiration was observed. However, photosystem II tolerance to extreme-heat damage (T50 ) was reduced from 50.0 ± 0.3°C to 48.5 ± 0.3°C under drought. Our results suggest that long-term reductions in precipitation, as projected across much of Amazonia by climate models, are unlikely to greatly alter the response of tropical forests to rising mean temperatures but may increase the risk of leaf thermal damage during heatwaves.

Complex Interactions Between Sex and Stress on Heroin Seeking.

Rationale: Stress plays a dual role in substance use disorders as a precursor to drug intake and a relapse precipitant. With heroin use at epidemic proportions in the United States, understanding interactions between stress disorders and opioid use disorder is vital and will aid in treatment of these frequently comorbid conditions. Objectives: Here, we combine assays of stress and contingent heroin self-administration (SA) to study behavioral adaptations in response to stress and heroin associated cues in male and female rats. Methods: Rats underwent acute restraint stress paired with an odor stimulus and heroin SA for subsequent analysis of stress and heroin cue reactivity. Lofexidine was administered during heroin SA and reinstatement testing to evaluate its therapeutic potential. Rats also underwent tests on the elevated plus maze, locomotor activity in a novel environment, and object recognition memory following stress and/or heroin. Results: A history of stress and heroin resulted in disrupted behavior on multiple levels. Stress rats avoided the stress conditioned stimulus and reinstated heroin seeking in response to it, with males reinstating to a greater extent than females. Lofexidine decreased heroin intake, reinstatement, and motor activity. Previous heroin exposure increased time spent in the closed arms of an elevated plus maze, activity in a round novel field, and resulted in object recognition memory deficits. Discussion: These studies report that a history of stress and heroin results in maladaptive coping strategies and suggests a need for future studies seeking to understand circuits recruited in this pathology and eventually help develop therapeutic approaches.

Fight hard or die trying: when plants face pathogens under heat stress.

In their natural environment, plants are exposed to biotic or abiotic stresses that occur sequentially or simultaneously. Plant responses to these stresses have been studied widely and have been well characterised in simplified systems involving single plant species facing individual stress. Temperature elevation is a major abiotic driver of climate change and scenarios have predicted an increase in the number and severity of epidemics. In this context, here we review the available data on the effect of heat stress on plant-pathogen interactions. Considering 45 studies performed on model or crop species, we discuss the possible implications of the optimum growth temperature of plant hosts and pathogens, mode of stress application and temperature variation on resistance modulations. Alarmingly, most identified resistances are altered under temperature elevation, regardless of the plant and pathogen species. Therefore, we have listed current knowledge on heat-dependent plant immune mechanisms and pathogen thermosensory processes, mainly studied in animals and human pathogens, that could help to understand the outcome of plant-pathogen interactions under elevated temperatures. Based on a general overview of the mechanisms involved in plant responses to pathogens, and integrating multiple interactions with the biotic environment, we provide recommendations to optimise plant disease resistance under heat stress and to identify thermotolerant resistance mechanisms.

Interactions of plant growth responses to spring freezing and summer drought: a multispecies comparison.

PREMISE OF THE STUDY: Freezing and drought both result in cellular dehydration, and similar physiological responses to these stressors may result in cross acclimation, whereby prior freezing exposure increases subsequent drought tolerance. We examined how spring freezing influences summer drought tolerance for a range of herbaceous old field species: 6 graminoids (Agrostis stolonifera, Arrhenatherum elatius, Bromus inermis, Festuca rubra, Lolium perenne, Poa compressa) and 2 forbs (Plantago lanceolata, Securigera varia), with the goal of examining the generality of cross acclimation responses. METHODS: We exposed the plants to -5°C in the spring and to a 3-week summer drought, and harvested the plants after a 3-week watering/recovery period. We also measured leaf soluble proteins and sugars to explore the potential mechanisms before and during drought stress. KEY RESULTS: For Agrostis stolonifera, Bromus inermis, Lolium perenne, Plantago lanceolata, and Poa compressa there was evidence of cross acclimation based on aboveground or belowground biomass, with a reduction in the severity of the drought effect for the plants previously exposed to freezing. Freezing and drought effects were additive for Arrhenatherum elatius, and for the remaining two species the test of the freezing-drought interaction was inconclusive, because significant drought and freezing effects did not co-occur. When present, freezing-drought interactions were not correlated with changes in leaf soluble protein or sugars. CONCLUSIONS: Our results reveal that the phenomenon of freezing-drought cross acclimation appears to be common in herbaceous species, and variation among species in cross acclimation indicates that multiple stresses could alter relative species abundances in plant communities.

The Effects of Combined Abiotic and Pathogen Stress in Plants: Insights From Salinity and Pseudomonas syringae pv lachrymans Interaction in Cucumber.

Plants are often challenged by abiotic and biotic stresses acting in combination and the response to combinatorial stress differs from that triggered by each factor individually. Although salinity and pathogens are major stressors limiting plant growth and productivity worldwide, their interaction is poorly understood. The reactions to pathogens overlap with those to abiotic stresses, and reactive oxygen species (ROS) and stress hormones represent central nodes in the interacting signaling pathways. Usually, abiotic stress negatively affects plant susceptibility to disease. Specific focus of this review is on cucumber plants exposed to salt stress and thereafter infected with Pseudomonas syringae pv lachrymans (Psl). We addressed this problem by discussing the changes in photochemistry, the antioxidant system, primary carbon metabolism, salicylic acid (SA) and abscisic acid (ABA) contents. Salt-treated plants were more prone to infection and this effect was determined by changes in the hormonal and redox balance as well as the carboxylate metabolism and activities of some NADPH-generating enzymes. Our detailed understanding of the interactive effects of biotic and abiotic stresses is fundamental to achieve enhanced tolerance to combination stress in agronomically important crops.

Genome-wide regulatory deterioration impedes adaptive responses to stress in inbred populations of Drosophila melanogaster.

Inbreeding depression is often intensified under environmental stress (i.e., inbreeding-stress interaction). Although the fitness consequences of this phenomenon are well-described, underlying mechanisms such as an increased expression of deleterious alleles under stress, or a lower capacity for adaptive responses to stress with inbreeding, have rarely been investigated. We investigated a fitness component (egg-to-adult viability) and gene-expression patterns using RNA-seq analyses in noninbred control lines and in inbred lines of Drosophila melanogaster exposed to benign temperature or heat stress. We find little support for an increase in the cumulative expression of deleterious alleles under stress. Instead, inbred individuals had a reduced ability to induce an adaptive gene regulatory stress response compared to controls. The decrease in egg-to-adult viability due to stress was most pronounced in the lines with the largest deviation in the adaptive stress response (R2 = 0.48). Thus, we find strong evidence for a lower capacity of inbred individuals to respond by gene regulation to stress and that this is the main driver of inbreeding-stress interactions. In comparison, the altered gene expression due to inbreeding at benign temperature showed no correlation with fitness and was pronounced in genomic regions experiencing the highest increase in homozygosity.

Salinity stress accelerates the effect of cadmium toxicity on soil N dynamics and cycling: Does joint effect of these stresses matter?

The objective of this study was to determine responses of soil nitrogen (N) transformation, microbial biomass N, and urease activity to the combined effect of cadmium (Cd) toxicity (0 and 30 mg kg-1) and NaCl stress (0, 7.5 and 15 dS m-1) in a clay loam soil unamended (0%) or amended with alfalfa residues (1%, w/w). Cd, NaCl, and alfalfa residues were added to the soil, and the mixtures were incubated for 90 days under standard laboratory conditions (25 ±â€¯1 °C and 70% of water holding capacity [WHC]). The results showed that salinity increased soil Cd availability and toxicity and subsequently decreased soil microbial N transformations (i.e., potential ammonification and nitrification as well as net N mineralization), arginine ammonification and nitrification rates, microbial biomass N, and urease activity. The adverse effects of salinity on soil microbial properties were greater in Cd-polluted than unpolluted soils, at high than low salinity levels, but were lower in residue-amended than unamended soils. These effects were mainly attributed to the increased Cd availability under saline conditions or the decreased Cd availability with residue addition. All the measured soil microbial attributes showed a negative correlation with the available Cd content in the soil. The interaction or combined effects of Cd and NaCl on soil microbial attributes were mostly synergistic in residue-unamended soils but antagonistic in residue-amended soils. The addition of organic residues to Cd-polluted soils may moderate salinity effect, and thus could stimulate the activity of ammonifiers and nitrifiers, as well as urease.

Primary carbon metabolism-related changes in cucumber exposed to single and sequential treatments with salt stress and bacterial infection.

This study examines how salt stress interacts with bacterial infection at the metabolic level. We measured chlorophyll a fluorescence as well as profiles of phosphoenolpyruvate carboxylase (PEPC), NADP-malic enzyme (NADP-ME), NADP-isocitrate dehydrogenase (NADP-ICDH) and fumarase activities, malic and citric acids contents and the expression of NADP-ICDH and NADP-ME in the organ-dependent (root vs leaves) response of cucumber plants exposed to individual or sequential action of salt stress (50 mM or 100 mM NaCl) and Pseudomonas syringae pv lachrymans (Psl). NaCl treatment, Psl infection and the combination of these stresses caused disturbances in the activity of photosystem II which were suggested to specifically transmit the stress signals. PEPC and NADP-ME were induced in cucumber plants under stress, confirming that in C3 plants they function in defence responses. The profiles of malate and citrate contents, PEPC as well as NADP-ICDH and NADP-ME activities and gene expression in response to a combination of salt and pathogen stresses differed from those provoked by individual stress with respect to the direction, intensity and timing. The results indicated that the most pronounced defence response related to the readjustment of the carbon metabolism was observed in the leaves of plants exposed to combined stress. Intense activity changes of NADPH-generating enzymes, NADP-ICDH and NADP-ME, characterized the tailored response to combined stress and could be important for the integration of defence mechanisms between organs.

Responses of African Grasses in the Genus Sporobolus to Defoliation and Sodium Stress: Tradeoffs, Cross-Tolerance, or Independent Responses?

In the Serengeti ecosystem of East Africa, grazing ungulates prefer areas with elevated grass Na, suggesting that some grasses tolerate both high soil Na and defoliation. We performed a factorial Na-by-defoliation greenhouse study with five abundant Sporobolus congeners to explore whether Serengeti grasses possess traits which: (i) confer tolerance to both Na and defoliation (cross-tolerance); (ii) display a tradeoff; or (iii) act independently in their tolerances. Our expectation was that related grasses would exhibit cross-tolerance when simultaneously subjected to Na and defoliation. Instead, we found that physiological tolerances and growth responses to Na and defoliation did not correlate but instead acted independently: species characterized by intense grazing in the field showed no growth or photosynthetic compensation for combined Na and defoliation. Additionally, in all but the highest Na dosage, mortality was higher when species were exposed to both Na and defoliation together. Across species, mortality rates were greater in short-statured species which occur on sodic soils in heavily grazed areas. Mortality among species was positively correlated with specific leaf area, specific root length, and relative growth rate, suggesting that rapidly growing species which invest in low cost tissues have higher rates of mortality when exposed to multiple stressors. We speculate that the prevalence of these species in areas of high Na and disturbance is explained by alternative strategies, such as high fecundity, a wide range of germination conditions, or further dispersal, to compensate for the lack of additional tolerance mechanisms.

Effects of nitrogen on the response of loblolly pine to drought: II. Biomass allocation and C : N balance.

Effects of drought on carbon and nitrogen dynamics and biomass allocation patterns of loblolly pine (Pinus taeda L.) seedlings grown under 4 different nitrogen levels were determined Nitrogen stress caused an increase in root-shoot ratio. increased Foliar and mot Starch concentration, decreased soluble sugar concentration, and decreased free amino acid concentration in well-watered seedlings. Drought cent-rally caused an increase in foliar hexoses and decreases in foliar find root starch and root hexoses. while not affecting foliar free amino acids. However, nitrogen stress arid drought interacted in affecting carbohydrate pools and seedling morphology. Foliar starch reserves in high-N seedling were rapidly metabolized under drought, white N-deficient seedling maintained similar starch concentrations throughout the drought stress cycle. After one drought cycle, the N-induced difference in root:shoot ratio was no longer significant. These results suggest that N-stressed seedlings partition a higher percentage of fixed carbon to storage, and that (his storage pool is less available For immediate Use than in seedlings grown under adequate N.