The number and intensity of heat waves has increased, and this trend is likely to continue throughout the 21st century. Often, heat waves are accompanied by drought conditions. It is projected that the global land area experiencing heat waves will double by 2020, and quadruple by 2040. Extreme heat events can impact a wide variety of tree functions. At the leaf level, photosynthesis is reduced, photooxidative stress increases, leaves abscise and the growth rate of remaining leaves decreases. In some species, stomatal conductance increases at high temperatures, which may be a mechanism for leaf cooling. At the whole plant level, heat stress can decrease growth and shift biomass allocation. When drought stress accompanies heat waves, the negative effects of heat stress are exacerbated and can lead to tree mortality. However, some species exhibit remarkable tolerance to thermal stress. Responses include changes that minimize stress on photosynthesis and reductions in dark respiration. Although there have been few studies to date, there is evidence of within-species genetic variation in thermal tolerance, which could be important to exploit in production forestry systems. Understanding the mechanisms of differing tree responses to extreme temperature events may be critically important for understanding how tree species will be affected by climate change.

Summary666I.Introduction667II.Biosynthesis667III.Meta‐analysis669IV.The type of stress influences the total amount of GLVs released669V.Herbivores can modulate the wound‐induced release of GLVs669VI.Fungal infection greatly induces GLV production672VII.Monocots and eudicots respond differentially to different types of stress673VIII.The type of stress does not influence the proportion of GLVs per chemical class673IX.The type of stress does influence the isomeric ratio within each chemical class674X.GLVs: from signal perception to signal transduction676XI.GLVs influence the C/N metabolism677XII.Interaction with plant hormones678XIII.General conclusions and unanswered questions678Acknowledgements679References679 Summary Plants respond to stress by releasing biogenic volatile organic compounds (BVOCs). Green leaf volatiles (GLVs), which are abundantly produced across the plant kingdom, comprise an important group within the BVOCs. They can repel or attract herbivores and their natural enemies; and they can induce plant defences or prime plants for enhanced defence against herbivores and pathogens and can have direct toxic effects on bacteria and fungi. Unlike other volatiles, GLVs are released almost instantly upon mechanical damage and (a)biotic stress and could thus function as an immediate and informative signal for many organisms in the plant's environment. We used a meta‐analysis approach in which data from the literature on GLV production during biotic stress responses were compiled and interpreted. We identified that different types of attackers and feeding styles add a degree of complexity to the amount of emitted GLVs, compared with wounding alone. This meta‐analysis illustrates that there is less variation in the GLV profile than we presumed, that pathogens induce more GLVs than insects and wounding, and that there are clear differences in GLV emission between monocots and dicots. Besides the meta‐analysis, this review provides an update on recent insights into the perception and signalling of GLVs in plants.

Priming refers to a mechanism whereby plants are sensitized to respond faster and/or more strongly to future pathogen attack. Here, we demonstrate that preexposure to the green leaf volatile Z-3-hexenyl acetate (Z-3-HAC) primed wheat (Triticum aestivum) for enhanced defense against subsequent infection with the hemibiotrophic fungus Fusarium graminearum. Bioassays showed that, after priming with Z-3-HAC, wheat ears accumulated up to 40% fewer necrotic spikelets. Furthermore, leaves of seedlings showed significantly smaller necrotic lesions compared with nonprimed plants, coinciding with strongly reduced fungal growth in planta. Additionally, we found that F. graminearum produced more deoxynivalenol, a mycotoxin, in the primed treatment. Expression analysis of salicylic acid (SA) and jasmonic acid (JA) biosynthesis genes and exogenous methyl salicylate and methyl jasmonate applications showed that plant defense against F. graminearum is sequentially regulated by SA and JA during the early and later stages of infection, respectively. Interestingly, analysis of the effect of Z-3-HAC pretreatment on SA-and JA-responsive gene expression in hormone-treated and pathogen-inoculated seedlings revealed that Z-3-HAC boosts JA-dependent defenses during the necrotrophic infection stage of F. graminearum but suppresses SA-regulated defense during its biotrophic phase. Together, these findings highlight the importance of temporally separated hormone changes in molding plant health and disease and support a scenario whereby the green leaf volatile Z-3-HAC protects wheat against Fusarium head blight by priming for enhanced JA-dependent defenses during the necrotrophic stages of infection.

The frequency and intensity of heat waves are predicted to increase. This study investigates whether heat waves would have the same impact as a constant increase in temperature with the same heat sum, and whether there would be any interactive effects of elevated [CO2 ] and soil moisture content. We grew Quercus rubra seedlings in treatment chambers maintained at either ambient or elevated [CO2 ] (380 or 700 μmol CO2 mol(-1) ) with temperature treatments of ambient, ambient +3 °C, moderate heat wave (+6 °C every other week) or severe heat wave (+12 °C every fourth week) temperatures. Averaged over a 4-week period, and the entire growing season, the three elevated temperature treatments had the same average temperature and heat sum. Half the seedlings were watered to a soil water content near field capacity, half to about 50% of this value. Foliar gas exchange measurements were performed morning and afternoon (9:00 and 15:00 hours) before, during and after an applied heat wave in August 2010. Biomass accumulation was measured after five heat wave cycles. Under ambient [CO2 ] and well-watered conditions, biomass accumulation was highest in the +3 °C treatment, intermediate in the +6 °C heat wave and lowest in the +12 °C heat wave treatment. This response was mitigated by elevated [CO2 ]. Low soil moisture significantly decreased net photosynthesis (Anet ) and biomass in all [CO2 ] and temperature treatments. The +12 °C heat wave reduced afternoon Anet by 23% in ambient [CO2 ]. Although this reduction was relatively greater under elevated [CO2 ], Anet values during this heat wave were still 34% higher than under ambient [CO2 ]. We concluded that heat waves affected biomass growth differently than the same amount of heat applied uniformly over the growing season, and that the plant response to heat waves also depends on [CO2 ] and soil moisture conditions.

SummaryHere, we investigated the effect of different heat-wave intensities applied at two atmospheric CO 2 concentrations ([CO 2 ]) on seedlings of two tree species, loblolly pine (Pinus taeda) and northern red oak (Quercus rubra).Seedlings were assigned to treatment combinations of two levels of [CO 2 ] (380 or 700 lmol mol À1 ) and four levels of air temperature (ambient, ambient +3°C, or 7-d heat waves consisting of a biweekly +6°C heat wave, or a monthly +12°C heat wave). Treatments were maintained throughout the growing season, thus receiving equal heat sums. We measured gas exchange and fluorescence parameters before, during and after a mid-summer heat wave. The +12°C heat wave, significantly reduced net photosynthesis (A net ) in both species and [CO 2 ] treatments but this effect was diminished in elevated [CO 2 ]. The decrease in A net was accompanied by a decrease in F v ′/F m ′ in P. taeda and Φ PSII in Q. rubra.Our findings suggest that, if soil moisture is adequate, trees will experience negative effects in photosynthetic performance only with the occurrence of extreme heat waves. As elevated [CO 2 ] diminished these negative effects, the future climate may not be as detrimental to plant communities as previously assumed.