Extreme summer heat waves are known to induce foliar and stem mortality in temperate forest ecosystems, yet our mechanistic knowledge of physiological thresholds for damage is lacking. Current spatiotemporal simulations of forest growth responses to climate change fail to explain the variability between co-occurring tree species to climate extremes, indicating a need for new model frameworks that include mechanistic understanding of trait-specific responses. In this context, using manipulative heat wave (hw) experiments we investigated ecophysiological responses and physiological recovery in four co-occurring temperate tree species of the southeastern United States including three deciduous angiosperms: southern red oak (Quercus falcata Michx.), shumard oak (Q. shumardii Buckl.) and, tulip-poplar (Liriodendron tulipifera L.) and one evergreen conifer: eastern white pine (Pinus strobus L.). The objectives were to investigate inter-specific differences in ecophysiological responses to hw events to understand mechanistic differences in resilience that may be useful for future model development. Two-year-old, well-irrigated potted saplings were exposed to progressively increasing extreme hw diurnal cycles followed by a recovery cycle, with peak midday air temperature increasing from 37 • C to a maximum of 51 • C on the third day of the hw. Plants were assessed for various photosynthetic and water use responses, chlorophyll fluorescence and photosystem-II (PSII) activity, leaf temperature and foliar pigments. Intense heat caused progressive down-regulation in net photosynthesis, but the stomata remained operational, which helped cool leaves through loss of latent heat. Even though whole plant transpiration increased for all species, the rate plateaued at higher hw events that allowed leaf temperature to exceed 45 • C, well beyond the optimal range. A significant increase in non-photochemical quenching over the hw cycles was evident in all species though indications of both transient and chronic PSII damage were evident in the most heat sensitive species, pine and tulip poplar. The oaks, especially Q. falcata, showed greater thermotolerance than other species with a higher threshold for photodamage to PSII, rapid overnight recovery of photoinhibition and minimal heat-induced canopy necrosis. We conclude that these co-occurring tree species exhibit large variability in thermotolerance and in their capability to repair both transient and chronic photodamage. Our results indicate that extreme heat induced damage to PSII within the leaf chloroplasts may be a mechanistic trait that can be used to project how different species respond to extreme weather events.

The induction and relaxation of photochemistry and non-photochemical quenching (NPQ) are not instantaneous and require time to respond to fluctuating environments. There is a lack of integrated understanding on how photochemistry and NPQ influence photosynthesis in fluctuating environments. We measured the induction and relaxation of chlorophyll a fluorescence and gas exchange in poplar and cotton at varying temperatures under saturating and fluctuating lights. When the light shifted from dark to high, the fraction of open reaction centers in photosystem II (qL) gradually increased while NPQ increased suddenly and then remained stable. Temperature significantly changed the response of qL but not that of NPQ during the dark to high light transition. Increased qL led to higher photosynthesis but their precise relationship was affected by NPQ and temperature. qL was significantly related to biochemical capacity. Thus, qL appears to be a strong indicator of the activation of carboxylase, leading to the similar dynamics between qL and photosynthesis. When the light shifted from high to low intensity, NPQ is still engaged at a high level, causing a stronger decline in photosynthesis. Our finding suggests that the dynamic effects of photochemistry and NPQ on photosynthesis depend on the phases of environmental fluctuations and interactive effects of light and temperature. Across the full spectra of light fluctuation, the slow induction of qL is a more important limiting factor than the slow relaxation of NPQ for photosynthesis in typical ranges of temperature for photosynthesis. The findings provided a new perspective to improve photosynthetic productivity with molecular biology under natural fluctuating environments.

Ward et al. Shrub Responses to WEW and eCO 2 decrease in R D25 for R. groenlandicum and in A ST for C. calyculata with increasing WEW, as well as an increase of A GR with eCO 2 in both species. Responses in newly emerged and overwintered leaves may reflect physiological acclimation or phenological changes in response to treatments.

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