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The Oak Ridge National Laboratory is planning to build the Second Target Station (STS) at the Spallation Neutron Source (SNS). STS will host a suite of novel instruments that complement the First Target Station’s beamline capabilities by offering an increased flux for cold neutrons and a broader wavelength bandwidth. A novel neutron imaging beamline, named the Complex, Unique, and Powerful Imaging Instrument for Dynamics (CUPI2D), is among the first eight instruments that will be commissioned at STS as part of the construction project. CUPI2D is designed for a broad range of neutron imaging scientific applications, such as energy storage and conversion (batteries and fuel cells), materials science and engineering (additive manufacturing, superalloys, and archaeometry), nuclear materials (novel cladding materials, nuclear fuel, and moderators), cementitious materials, biology/medical/dental applications (regenerative medicine and cancer), and life sciences (plant–soil interactions and nutrient dynamics). The innovation of this instrument lies in the utilization of a high flux of wavelength-separated cold neutrons to perform real time in situ neutron grating interferometry and Bragg edge imaging—with a wavelength resolution of δλ/λ ≈ 0.3%—simultaneously when required, across a broad range of length and time scales. This manuscript briefly describes the science enabled at CUPI2D based on its unique capabilities. The preliminary beamline performance, a design concept, and future development requirements are also presented.

Heat and drought affect plant chemical defenses and thereby plant susceptibility to pests and pathogens. Monoterpenes are of particular importance for conifers as they play critical roles in defense against bark beetles. To date, work seeking to understand the impacts of heat and drought on monoterpenes has primarily focused on young potted seedlings, leaving it unclear how older age classes that are more vulnerable to bark beetles might respond to stress. Furthermore, we lack a clear picture of what carbon resources might be prioritized to support monoterpene synthesis under drought stress. To address this, we measured needle and woody tissue monoterpene concentrations and physiological variables simultaneously from mature piñon pines (Pinus edulis) from a unique temperature and drought manipulation field experiment. While heat had no effect on total monoterpene concentrations, trees under combined heat and drought stress exhibited ~ 85% and 35% increases in needle and woody tissue, respectively, over multiple years. Plant physiological variables like maximum photosynthesis each explained less than 10% of the variation in total monoterpenes for both tissue types while starch and glucose + fructose measured 1-month prior explained ~ 45% and 60% of the variation in woody tissue total monoterpene concentrations. Although total monoterpenes increased under combined stress, some key monoterpenes with known roles in bark beetle ecology decreased. These shifts may make trees more favorable for bark beetle attack rather than well defended, which one might conclude if only considering total monoterpene concentrations. Our results point to cumulative and synergistic effects of heat and drought that may reprioritize carbon allocation of specific non-structural carbohydrates toward defense. Supplementary Information The online version contains supplementary material available at 10.1007/s00442-021-05058-8.

Understanding and predicting the relationship between leaf temperature ( T leaf ) and air temperature ( T air ) is essential for projecting responses to a warming climate, as studies suggest that many forests are near thermal thresholds for carbon uptake. Based on leaf measurements, the limited leaf homeothermy hypothesis argues that daytime T leaf is maintained near photosynthetic temperature optima and below damaging temperature thresholds. Specifically, leaves should cool below T air at higher temperatures (i.e., > ∼25–30°C) leading to slopes T leaf / T air relationships and substantial carbon uptake when leaves are cooler than air. This hypothesis implies that climate warming will be mitigated by a compensatory leaf cooling response. A key uncertainty is understanding whether such thermoregulatory behavior occurs in natural forest canopies. We present an unprecedented set of growing season canopy-level leaf temperature ( T can ) data measured with thermal imaging at multiple well-instrumented forest sites in North and Central America. Our data do not support the limited homeothermy hypothesis: canopy leaves are warmer than air during most of the day and only cool below air in mid to late afternoon, leading to T can / T air slopes >1 and hysteretic behavior. We find that the majority of ecosystem photosynthesis occurs when canopy leaves are warmer than air. Using energy balance and physiological modeling, we show that key leaf traits influence leaf-air coupling and ultimately the T can / T air relationship. Canopy structure also plays an important role in T can dynamics. Future climate warming is likely to lead to even greater T can , with attendant impacts on forest carbon cycling and mortality risk.

The phloem provides a pathway for products of photosynthesis to be transported to different parts of the plant for consumption or storage. The Münch pressure flow hypothesis (PFH) is considered the leading framework to mathematically represent this transport. It assumes that osmosis provides the necessary pressure differences to drive the fluid flow and sucrose within the phloem. Mathematical models utilizing the PFH approximate the phloem by a relatively rigid semi-permeable tube. However, the phloem consists of living cells that contract and expand in response to pressure fluctuations. The effect of membrane elasticity on osmotically driven sucrose front speed has rarely been considered and frames the scope here. Laboratory experiments were conducted to elucidate the elastic-to-plastic pressure-deformation relation in membranes and their effect on sucrose front speeds. It is demonstrated that membrane elasticity acts to retard the sucrose front speed. The retardation emerges because some of the osmotic pressure performs mechanical work to expand the membrane instead of pressurizing water. These results offer a novel perspective about the much discussed presence of sieve plates through-out the phloem acting as structural dampers.

In Arctic ecosystems, increasing temperatures are driving the expansion of nitrogen (N) fixing shrubs across tundra landscapes. The implications of this expansion to the biogeochemistry of Arctic ecosystems are of critical importance and more work is needed to better understand the form, availability, and transportation potential of N from these shrubs across a variety of Arctic landscapes. To gain insights into the processes controlling N within a permafrost hillslope system, the spatiotemporal variability of nitrate (NO3-) and its environmental controls were investigated at an alder (Alnus viridis spp. fruticosa) dominated permafrost tundra landscape in the Seward Peninsula, Alaska, USA. Soil pore water was collected from locations within alder shrubland growing along a well-drained hillslope and was compared to soil pore water collected from locations outside (upslope, downslope, and between) the alder shrubland. Soil pore water collected within alder shrubland had an average NO3-N (nitrogen from nitrate) concentration of 4.27±8.02 mg L−1 and differed significantly from locations outside alder shrubland (0.23±0.83 mg L−1; p). Temporal variation in NO3-N within and downslope of alder shrubland co-occurred with precipitation events where NO3- that accumulated in the soil was likely flushed downslope during rainfall. These findings have important implications for nutrient availability and mobility in N-limited permafrost systems that are experiencing shrub expansion in response to a warming Arctic.

Understanding mass transport of photosynthates in the phloem of plants is necessary for predicting plant carbon allocation, productivity, and responses to water and thermal stress. Several hypotheses about optimization of phloem structure and function and limitations of phloem transport under drought have been proposed and tested with models and anatomical data. However, the true impact of radial water exchange of phloem conduits with their surroundings on mass transport of photosynthates has not been addressed. Here, the physics of the Munch mechanism of sugar transport is re-evaluated to include local variations in viscosity resulting from the radial water exchange in two dimensions (axial and radial) using transient flow simulations. Model results show an increase in radial water exchange due to a decrease in sap viscosity leading to increased sugar front speed and axial mass transport across a wide range of phloem conduit lengths. This increase is around 40% for active loaders (e.g. crops) and around 20% for passive loaders (e.g. trees). Thus, sugar transport operates more efficiently than predicted by previous models that ignore these two effects. A faster front speed leads to higher phloem resiliency under drought because more sugar can be transported with a smaller pressure gradient.

Terrestrial gross primary production (GPP) is the basis of vegetation growth and food production globally1 and plays a critical role in regulating atmospheric CO2 through its impact on ecosystem carbon balance. Even though higher CO2 concentrations in future decades can increase GPP2, low soil water availability, heat stress and disturbances associated with droughts could reduce the benefits of such CO2 fertilization. Here we analysed outputs of 13 Earth system models to show an increasingly stronger impact on GPP by extreme droughts than by mild and moderate droughts over the twenty-first century. Due to a dramatic increase in the frequency of extreme droughts, the magnitude of globally averaged reductions in GPP associated with extreme droughts was projected to be nearly tripled by the last quarter of this century (2075–2099) relative to that of the historical period (1850–1999) under both high and intermediate GHG emission scenarios. By contrast, the magnitude of GPP reductions associated with mild and moderate droughts was not projected to increase substantially. Our analysis indicates a high risk of extreme droughts to the global carbon cycle with atmospheric warming; however, this risk can be potentially mitigated by positive anomalies of GPP associated with favourable environmental conditions. Terrestrial primary productivity will increase with CO2 fertilization, but water limitation will decrease this positive effect. Analyses of Earth system model projections show that extreme droughts will have a much stronger impact on future productivity than mild and moderate droughts.

Plant survival during environmental stress greatly affects ecosystem carbon (C) cycling, and plant-microbe interactions are central to plant stress survival. The release of C-rich root exudates is a key mechanism plants use to manage their microbiome, attracting beneficial microbes and/or suppressing harmful microbes to help plants withstand environmental stress. However, a critical knowledge gap is how plants alter root exudate concentration and composition under varying stress levels. In a greenhouse study, we imposed three drought treatments (control, mild, severe) on blue grama (Bouteloua gracilis Kunth Lag. Ex Griffiths), and measured plant physiology and root exudate concentration and composition using GC-MS, NMR, and FTICR. With increasing drought severity, root exudate total C and organic C increased concurrently with declining predawn leaf water potential and photosynthesis. Root exudate composition mirrored the physiological gradient of drought severity treatments. Specific compounds that are known to alter plant drought responses and the rhizosphere microbiome mirrored the drought severity-induced root exudate compositional gradient. Despite reducing C uptake, these plants actively invested C to root exudates with increasing drought severity. Patterns of plant physiology and root exudate concentration and composition co-varied along a gradient of drought severity.

Mass transport of photosynthates in the phloem of plants is necessary for describing plant carbon allocation, productivity, and responses to water and thermal stress. Several hypotheses about optimization of phloem structure and function and limitations of phloem transport under drought have been proposed, and tested with models and anatomical data. However, the true impact of radial water exchange of phloem conduits with their surroundings on mass transport of photosynthates has not been addressed. Here, the physics of the Munch mechanism of sugar transport is re-evaluated to include local variations in viscosity resulting from the radial water exchange in two dimensions (axial and radial). Model results show that radial water exchange pushes sucrose away from conduit walls thereby reducing wall frictional stress due to a decrease in sap viscosity and an increase in sugar concentration in the central region of the conduit. These two co-occurring effects lead to increased sugar front speed and axial mass transport across a wide range of phloem conduit lengths. Thus, sugar transport operates more efficiently than predicted by previous models that ignore these two effects. A faster front speed leads to higher phloem resiliency under drought because more sugar can be transported with a smaller pressure gradient.SummaryThe overall speed of sap increased by including a concentrationdependent viscosity in axial and radial directions.

Short-term plant respiration (R) increases exponentially with rising temperature, but drought could reduce respiration by reducing growth and metabolism. Acclimation may alter these responses. We examined if species with different drought responses would differ in foliar R response to +4.8°C temperature and -45% precipitation in a field experiment with mature pinon and juniper trees, and if any differences between species were related to differences in photosynthesis rates, shoot growth and non-structural carbohydrates (NSCs). Short-term foliar R had a Q10 of 1.6 for pinon and 2.6 for juniper. Pinon foliar R did not respond to the +4.8°C temperatures, but R increased 1.4x for juniper. Across treatments, pinon foliage had higher growth, lower NSC content, 29% lower photosynthesis rates, and 44% lower R than juniper. Removing 45% precipitation had little impact on R for either species. Species differences in the response of R under elevated temperature were related to substrate availability and stomatal response to leaf water potential. Despite not acclimating to the higher temperature and having higher R than pinon, greater substrate availability in juniper suggests it could supply respiratory demand for much longer than pinon. Species responses will be critical in ecosystem response to a warmer climate. This article is protected by copyright. All rights reserved.

In Arctic ecosystems, increasing temperatures are driving the expansion of nitrogen (N) fixing shrubs across tundra landscapes. The implications of this expansion to the biogeochemistry of Arctic ecosystems is of critical importance, yet many details about the form, location, and availability of N from these shrubs remain unknown. To address this knowledge gap, the spatiotemporal variability of nitrate (NO3−) and its environmental and edaphic controls were investigated at an alder (Alnus viridis spp. fruticosa) dominated permafrost tundra landscape in the Seward Peninsula, Alaska, USA. Soil pore water was collected from locations within alder shrubland growing along a well-drained hillslope and compared to soil pore water collected from locations outside (upslope, downslope, and between) the alder shrubland. δ15N and δ18O of soil pore water were consistent with the predicted range of NO3− produced through microbial degradation of N-rich alder shrub organic matter. Soil pore water collected within alder shrubland had an average NO3− concentration of (4.27 ± 8.02 mg L−1) and differed significantly from locations outside alder shrubland (0.23 ± 0.83 mg L−1; p 3− within and downslope of alder shrubland corresponded to precipitation events, where NO3− accumulated in the soil was flushed downslope during rainfall. Enrichment of both δ15N and δ18O isotopes at wetter downslope locations indicate that denitrification buffered the mobility and spatial extent of NO3−. These findings have important implications for nutrient production and mobility in N-limited permafrost systems that are experiencing shrub expansion in response to a warming Arctic.

On-going climate change is increasing the risk of drought stress across large areas worldwide. Such drought events decrease ecosystem productivity and have been increasingly linked to tree mortality. Understanding how trees respond to water shortage is key to predicting the future of ecosystem functions. Phloem is at the core of the tree functions, moving resources such as non-structural carbohydrates, nutrients, and defence and information molecules across the whole plant. Phloem function and ability to transport resources is tightly controlled by the balance of carbon and water fluxes within the tree. As such, drought is expected to impact phloem function by decreasing the amount of available water and new photoassimilates. Yet, the effect of drought on the phloem has received surprisingly little attention in the last decades. Here we review existing knowledge on drought impacts on phloem transport from loading and unloading processes at cellular level to possible effects on long-distance transport and consequences to ecosystems via ecophysiological feedbacks. We also point to new research frontiers that need to be explored to improve our understanding of phloem function under drought. In particular, we show how phloem transport is affected differently by increasing drought intensity, from no response to a slowdown, and explore how severe drought might actually disrupt the phloem transport enough to threaten tree survival. Because transport of resources affects other organisms interacting with the tree, we also review the ecological consequences of phloem response to drought and especially predatory, mutualistic and competitive relations. Finally, as phloem is the main path for carbon from sources to sink, we show how drought can affect biogeochemical cycles through changes in phloem transport. Overall, existing knowledge is consistent with the hypotheses that phloem response to drought matters for understanding tree and ecosystem function. However, future research on a large range of species and ecosystems is urgently needed to gain a comprehensive understanding of the question.

International audience; Sucrose is among the main products of photosynthesis that are deemed necessary for plant growth and survival. It is produced in the mesophyll cells of leaves and translocated to different parts of the plant through the phloem. Progress in understanding this transport process remains fraught with experimental difficulties, thereby prompting interest in theoretical approaches and laboratory studies. The Munch pressure and mass flow model is one of the accepted hypotheses describing the physics of sucrose transport in the phloem. It is based on osmosis creating an energy potential difference between the source and the sink. The flow responding to this energy potential is assumed laminar and described by the Hagen-Poiseuille equation. This study revisits such osmotically driven flows in tubes with membrane walls by including the effects of Taylor dispersion on mass transport. This effect has been overlooked in phloem flow studies. Taylor dispersion can increase the effective transport of solutes by increasing the apparent diffusion coefficient. It is shown that, in addition to the conventional diffusive correction derived for impermeable tube walls, a new adjustment to the mean advective terms arises because of osmotic effects. Because the molecular Schmidt number is very large for sucrose in water, the sucrose front speed and travel times have a direct dependence on the Peclet number for different ranges of the Munch number. This study establishes upper limits on expected Taylor dispersion enhancement of sucrose transport.

Leaf litter decomposition is a major carbon input to soil, making it a target for increasing soil carbon storage through microbiome engineering. We expand upon previous findings to show with multiple leaf litter types that microbial composition can drive variation in carbon flow from litter decomposition and specific microbial community features are associated with synonymous patterns of carbon flow among litter types. Although plant litter type selects for different decomposer communities, within a litter type, microbial composition drives variation in the quantity of dissolved organic carbon (DOC) measured at the end of the decomposition period. Bacterial richness was negatively correlated with DOC quantity, supporting our hypothesis that across multiple litter types there are common microbial traits linked to carbon flow patterns. Variation in DOC abundance (i.e. high versus low DOC) driven by microbial composition is tentatively due to differences in bacterial metabolism of labile compounds, rather than catabolism of non-labile substrates such as lignin. The temporal asynchrony of metabolic processes across litter types may be a substantial impediment to discovering more microbial features common to synonymous patterns of carbon flow among litters. Overall, our findings support the concept that carbon flow may be programmed by manipulating microbial community composition.

Changing climates can cause shifts in temperature and precipitation, resulting in warming and drought in some regions. Although each of these factors has been shown to detrimentally affect forest ecosystems worldwide, information on the impacts of the combined effects of warming and drought is lacking. Forest trees rely on mutualistic root-associated fungi that contribute significantly to plant health and protection against climate stresses. We used a six-year, ecosystem-scale temperature and precipitation manipulation experiment targeted to simulate the climate in 2100 in the Southwestern United States to quantify the effects of drought, warming and combined drought and warming on the root colonization (abundance), species composition and diversity of ectomycorrhizal fungi (EMF), and dark septate fungal endophytes in a widespread woodland tree, pinyon pine (Pinus edulis E.). Our results show that pinyon shoot growth after 6 years of these treatments was reduced more by drought than warming. The combined drought and warming treatment reduced the abundance and diversity of EMF more than either treatment alone. Individual ectomycorrhizal fungal taxa, including the drought tolerant Cenococcum geophilum, were present in all treatments but the combined drought and warming treatment. The combined drought and warming treatment also reduced the abundance of dark septate endophytes (DSE), but did not affect their diversity or species composition. The current year shoot growth of the trees correlated positively with ectomycorrhizal fungal diversity, highlighting the importance of diversity in mutualistic relationships to plant growth. Our results suggest that EMF may be more important than DSE to aboveground growth in P. edulis, but also more susceptible to the negative effects of combined climate stressors.

Examining factors that influence seedling establishment is essential for predicting the impacts of climate change on tree species’ distributions. Seedlings originating from contrasting climates differentially express functional traits related to water and nutrient uptake and drought resistance that reflect their climate of origin and influence their responses to drought. Soil microbes may improve seedling establishment because they can enhance water and nutrient uptake and drought resistance. However, the relative influence of soil microbes on the expression of these functional traits between seedling families or populations from contrasting climates is unknown. To determine if soil microbes may differentially alter functional traits to enhance water and nutrient uptake and drought resistance between dry and wet families, seeds of loblolly pine families from the driest and wettest ends of its geographic range (dry, wet) were planted in sterilized sand (controls) or in sterilized sand inoculated with a soil microbial community (inoculated). Functional traits related to seedling establishment (germination), water and nutrient uptake and C allocation (root:shoot biomass ratio, root exudate concentration, leaf C:N, leaf N isotope composition (δ15N)), and drought resistance (turgor loss point, leaf carbon isotope composition (δ13C)) were measured. Then, plants were exposed to a drought treatment and possible shifts in photosynthetic performance were monitored using chlorophyll fluorescence. Inoculated plants exhibited significantly greater germination than controls regardless of family. The inoculation treatment significantly increased root:shoot biomass ratio in the wet family but not in the dry family, suggesting soil microbes alter functional traits that improve water and nutrient uptake more so in a family originating from a wetter climate than in a family originating from a drier climate. Microbial effects on photosynthetic performance during drought also differed between families, as photosynthetic performance of the dry inoculated group declined fastest. Regardless of treatment, the dry family exhibited a greater root:shoot biomass ratio, root exudate concentration, and leaf δ15N than the wet family. This indicates that the dry family allocated more resources belowground than the wet and the two family may have used different sources of plant available N, which may be related to their contrasting climates of origin and influence their drought resistance. Examination of variation in impacts of soil microbes on seedling physiology improves efforts to enhance seedling establishment and beneficial plant-microbe interactions under climate change.

The leaf area to sapwood area ratios of trees (Al:AS) can shift to maintain homeostatic gas exchange per unit leaf area in response to climate variability. We tested the hypothesis that trees alter their Al:AS ratios in response to long-term warming and reduced precipitation in order to maintain leaf-specific gas exchange rates under more stressful conditions. Whole-tree Al:AS was measured on mature piñon pine (Pinus edulis Engelm.) and one-seed juniper (Juniperus monosperma (Engelm.) Sarg.) trees after 5 years (2012-16) of chronic exposure to increased temperature (+4.8 °C), precipitation reduction (-45%), or both simultaneously. No difference was found in Al:As among treatments for either species. Associated with this lack of shift in Al:As were large changes in pre-dawn leaf water potential and stomatal conductance, consistent with theoretical expectations of interactions between leaf and whole-tree hydraulic supply. Our results suggest that a lack of whole-tree acclimation in Al:As results in the reductions in plant gas exchange and water status associated with long-term warming and reduced precipitation in semi-arid woodlands.

We investigated stem radial growth and water storage dynamics of 2 conifer species differing in hydraulic carbon strategies, Juniperus monosperma and Pinus edulis, under conditions of ambient, drought (∼45% reduction in precipitation), heat (∼4.8 °C temperature increase), and the combination of drought + heat, in 2013 and 2014. Juniper maintained low growth across all treatments. Overall, the relatively isohydric pinon pine showed significantly greater growth and water storage recharge than the relatively anisohydric juniper across all treatments in the average climate year (2014) but no differences in the regionally dry year (2013). Pinon pine ceased growth at a constant predawn water potential across all treatments and at a less negative water potential threshold than juniper. Heat has a greater negative impact on pinon pines' growth and water storage than drought, whereas juniper was, in contrast, unaffected by heat but strongly impacted by drought. The whole-plant hydraulic carbon strategies, in this case captured using the isohydric/anisohydric concept, translate into alternative growth and water storage strategies under drought and heat conditions.

Drought impacts on phloem transport have attracted attention only recently, despite the well-established, and empirically verified theories on drought impacts on water transport in plants in general. This is because studying phloem transport is challenging. Phloem tissue is relatively small and delicate, and it has often been assumed not to be impacted by drought, or having insignificant impact on plant function or survival compared to the xylem. New evidence, however, suggests that drought responses of the phloem might hold the key for predicting plant survival time during drought or revival capacity after drought. This review summarizes current theories and empirical evidence on how drought might impact phloem transport, and evaluates these findings in relation to plant survival during drought.

Plants close their stomata during drought to avoid excessive water loss, but species differ in respect to the drought severity at which stomata close. The stomatal closure point is related to xylem anatomy and vulnerability to embolism, but it also has implications for phloem transport and possibly phloem anatomy to allow sugar transport at low water potentials. Desiccation-tolerant plants that close their stomata at severe drought should have smaller xylem conduits and/or fewer and smaller interconduit pits to reduce vulnerability to embolism but more phloem tissue and larger phloem conduits compared with plants that avoid desiccation. These anatomical differences could be expected to increase in response to long-term reduction in precipitation. To test these hypotheses, we used tridimensional synchroton X-ray microtomograph and light microscope imaging of combined xylem and phloem tissues of 2 coniferous species: one-seed juniper (Juniperus monosperma) and pinon pine (Pinus edulis) subjected to precipitation manipulation treatments. These species show different xylem vulnerability to embolism, contrasting desiccation tolerance, and stomatal closure points. Our results support the hypothesis that desiccation tolerant plants require higher phloem transport capacity than desiccation avoiding plants, but this can be gained through various anatomical adaptations in addition to changing conduit or tissue size.

Disentangling the relative impacts of precipitation reduction and vapour pressure deficit (VPD) on plant water dynamics and determining whether acclimation may influence these patterns in the future is an important challenge. Here, we report sap flux density (FD ), stomatal conductance (Gs ), hydraulic conductivity (KL ) and xylem anatomy in pinon pine (Pinus edulis) and juniper (Juniperus monosperma) trees subjected to five years of precipitation reduction, atmospheric warming (elevated VPD) and their combined effects. No acclimation occurred under precipitation reduction: lower Gs and FD were found for both species compared to ambient conditions. Warming reduced the sensibility of stomata to VPD for both species but resulted in the maintenance of Gs and FD to ambient levels only for pinon. For juniper, reduced soil moisture under warming negated benefits of stomatal adjustments and resulted in reduced FD , Gs and KL . Although reduced stomatal sensitivity to VPD also occurred under combined stresses, reductions in Gs , FD and KL took place to similar levels as under single stresses for both species. Our results show that stomatal conductance adjustments to high VPD could minimize but not entirely prevent additive effects of warming and drying on water use and carbon acquisition of trees in semi-arid regions.

© 2016 John Wiley & Sons Ltd Laboratory studies on artificial leaves suggest that leaf thermal dynamics are strongly influenced by the two-dimensional size and shape of leaves and associated boundary layer thickness. Hot environments are therefore said to favour selection for small, narrow or dissected leaves. Empirical evidence from real leaves under field conditions is scant and traditionally based on point measurements that do not capture spatial variation in heat load. We used thermal imagery under field conditions to measure the leaf thermal time constant (τ) in summer and the leaf-to-air temperature difference (∆T) and temperature range across laminae (Trange) during winter, autumn and summer for 68 Proteaceae species. We investigated the influence of leaf area and margin complexity relative to effective leaf width (we), the latter being a more direct indicator of boundary layer thickness. Normalized difference of margin complexity had no or weak effects on thermal dynamics, but we strongly predicted τ and ∆T, whereas leaf area influenced Trange. Unlike artificial leaves, however, spatial temperature distribution in large leaves appeared to be governed largely by structural variation. Therefore, we agree that small size, specifically we, has adaptive value in hot environments but not with the idea that thermal regulation is the primary evolutionary driver of leaf dissection.