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Satellite and tower-based metrics of forest-scale photosynthesis generally increase with dry season progression across central Amazônia, but the underlying mechanisms lack consensus. We conducted demographic surveys of leaf age composition, and measured the age dependence of leaf physiology in broadleaf canopy trees of abundant species at a central eastern Amazon site. Using a novel leaf-to-branch scaling approach, we used these data to independently test the much-debated hypothesis – arising from satellite and tower-based observations – that leaf phenology could explain the forest-scale pattern of dry season photosynthesis. Stomatal conductance and biochemical parameters of photosynthesis were higher for recently mature leaves than for old leaves. Most branches had multiple leaf age categories simultaneously present, and the number of recently mature leaves increased as the dry season progressed because old leaves were exchanged for new leaves. These findings provide the first direct field evidence that branch-scale photosynthetic capacity increases during the dry season, with a magnitude consistent with increases in ecosystem-scale photosynthetic capacity derived from flux towers. Interactions between leaf age-dependent physiology and shifting leaf age-demographic composition are sufficient to explain the dry season photosynthetic capacity pattern at this site, and should be considered in vegetation models of tropical evergreen forests.

Specialist herbivores and pathogens could induce negative conspecific density dependence among their hosts and thereby contribute to the diversity of plant communities. A small number of hyperdiverse genera comprise a large portion of tree diversity in tropical forests. These closely related congeners are likely to share natural enemies. Diverse defenses could still allow congeners to partition niche space defined by natural enemies, but interspecific differences in defenses would have to exceed intraspecific variation in defenses. We ask whether interspecific variation in secondary chemistry exceeds intraspecific variation for species from four hyperdiverse tropical tree genera. We used novel methods to quantify chemical structural similarity for all compounds present in methanol extracts of leaf tissue. We sought to maximize intraspecific variation by selecting conspecific leaves from different ontogenetic stages (expanding immature vs. fully hardened mature), different light environments (deep understory shade vs. large forest gaps), and different seasons (dry vs. wet). Chemical structural similarity differed with ontogeny, light environment, and season, but interspecific differences including those among congeneric species were much larger. Our results suggest that species differences in secondary chemistry are large relative to within-species variation, perhaps sufficiently large to permit niche segregation among congeneric tree species based on chemical defenses.

Premise The size and shape (physiognomy) of woody, dicotyledonous angiosperm leaves are correlated with climate. These relationships are the basis for multiple paleoclimate proxies. Here we test whether Vitis exhibits phenotypic plasticity and whether physiognomy varies along the vine. Methods We used Digital Leaf Physiognomy (DiLP) to measure leaf characters of four Vitis species from the USDA Germplasm Repository (Geneva, New York) from the 2012–2013 and 2014–2015 leaf‐growing seasons, which had different environmental conditions. Results Leaf shape changed allometrically through developmental stages; early stages were more linear than later stages. There were significant differences in physiognomy in the same developmental stage between the growing seasons, and species had significant differences in mean physiognomy between growing seasons. Phenotypic plasticity was defined as changes between growing seasons after controlling for developmental stage or after averaging all developmental stages. Vitis amurensis and V. riparia had the greatest phenotypic plasticity. North American species exhibited significant differences in tooth area:blade area. Intermediate developmental stages were most likely to exhibit phenotypic plasticity, and only V. amurensis exhibited phenotypic plasticity in later developmental stages. Conclusions Leaves have variable phenotypic plasticity along the vine. Environmental signal was strongest in intermediate developmental stages. This is significant for leaf physiognomic‐paleoclimate proxies because these leaves are likely the most common in leaf litter and reflect leaves primarily included in paleoclimate reconstructions. Early season and early developmental stages have the potential to be confounding factors but are unlikely to exert significant influence because of differential preservation potential.

IntroductionThe oxylipin methyl jasmonate (MeJA) is a plant hormone active in response signalling and defence against herbivores. Although MeJA is applied experimentally to mimic herbivory and induce plant defences, its downstream effects on the plant metabolome are largely uncharacterized, especially in the context of primary growth and tissue-specificity of the response.ObjectivesWe investigated the effects of MeJA-simulated and real caterpillar herbivory on the foliar metabolome of the wild plant Brassica nigra and monitored the herbivore-induced responses in relation to leaf ontogeny.MethodsAs single or multiple herbivory treatments, MeJA- and mock-sprayed plants were consecutively exposed to caterpillars or left untreated. Gas chromatography (GC) and liquid chromatography (LC) time-of-flight mass-spectrometry (TOF-MS) were combined to analyse foliar compounds, including central primary and specialized defensive plant metabolites.ResultsPlant responses were stronger in young leaves, which simultaneously induced higher chlorophyll levels. Both MeJA and caterpillar herbivory induced similar, but not identical, accumulation of tricarboxylic acids (TCAs), glucosinolates (GSLs) and phenylpropanoids (PPs), but only caterpillar feeding led to depletion of amino acids. MeJA followed by caterpillars caused higher induction of defence compounds, including a three-fold increase in the major defence compound allyl-GSL (sinigrin). When feeding on MeJA-treated plants, caterpillars gained less weight indicative of the reduced host-plant quality and enhanced resistance.ConclusionsThe metabolomics approach showed that plant responses induced by herbivory extend beyond the regulation of defence metabolism and are tightly modulated throughout leaf development. This leads to a new understanding of the plant metabolic potential that can be exploited for future plant protection strategies.

Summary Leaf photosynthetic properties, for example the maximum carboxylation velocity or Vc,max, change with leaf age due to ontogenetic processes. This study introduces an optimal dynamic allocation scheme to model changes in leaf‐level photosynthetic capacity as a function of leaf biochemical constraints (costs of synthesis and defence), nitrogen availability and other environmental factors (e.g. light). The model consists of a system of equations describing RuBisCO synthesis and degradation within chloroplasts, defence and ageing at leaf levels, nitrogen transfer and carbon budget at plant levels. Model results show that optimal allocation principles explained RuBisCO dynamics with leaf age. An approximated analytical solution can reproduce the basic pattern of RuBisCO and Vc,max in rice and in two tropical tree species. The model also reveals leaf life complementarities that remained unexplained in previous approaches, as the interplay between Vc,max at maturation, life span and the decline in photosynthetic capacity with age. Furthermore, it explores the role of defence, which is not implemented in current models. This framework covers some of the existing gaps in integrating multiple processes across plant organs (chloroplast, leaf and whole plant) and is a first‐step towards representing mechanistically leaf ontogenetic processes into physiological and ecosystem models.

Leaves, considered as the ‘source’ organs, depend on the development stages because of the age-dependent photosynthesis and assimilation of leaves. However, the molecular mechanisms of age-dependent limitations on the function of leaves are seldom reported. In the present study, the photosynthesis-related characteristics and photoassimilates were investigated in grape leaves at six different age groups (Ll to L6) at micro-morphological, biochemical, and molecular levels. These results showed lower expression levels of genes associated with stomatal development, and chl biosynthesis resulted in fewer stomata and lowered chlorophyll a/b contents in L1 when compared to L3 and L5. The DEGs between L5 and L3/L1 were largely distributed at stomatal movement, carbon fixation, and sucrose and starch metabolism pathways, such as STOMATAL ANION CHANNEL PROTEIN 1 (SLAC1), FRUCTOSE-1,6-BISPHOSPHATE ALDOLASE (FBA1), SUCROSE-PHOSPHATE SYNTHASE (SPP1), and SUCROSE-PHOSPHATE PHOSPHATASE (SPS2, 4). These genes could be major candidate genes leading to increased photosynthesis capacity and sugar content in L5. The accumulation of starch grains in the chloroplast and palisade tissue of L5 and higher transcription levels of genes related to starch biosynthesis in L5 further supported the high ability of L5 to produce photoassimilates. Hence, our results provide insights for understanding different photosynthetic functions in age-dependent leaves in grape plants at the molecular level.

Ultraviolet radiation (UV) represents a significant abiotic stress, affecting green plants. Phenolic compounds have been suggested as components involved in plant photoprotective adaptation. We used a unique combination of experimental (LED lighting and leaf tagging) and analytical (unbiased, or untargeted, metabolomics) approaches to study the effects of high (approximating mid-summer) and low (approximating winter) levels of UVA on the expression of phenolic compounds. These consisted of river red gum (Eucalyptus camaldulensis ssp. camaldulensis) of five provenances. The geographically separated provenances used in our study spanned the lowest and highest latitudes of the range of this subspecies. The concentrations of gallotannins and ellagitannins (i.e., hydrolysable tannins) increased most under high levels of UVA, but responses only differed slightly among provenances. The most substantial changes in the composition of phenolic compounds were associated with leaf age. Overall, 3-month-old (herein, termed ‘young’) leaves had substantially different phenolic compositions to 6- and 12-month-old (‘old’) leaves. Hydrolysable tannins were more abundant in young leaves, whereas pedunculagin, catechin, and kaempferol galloyl glucoses were more abundant in old leaves. High levels of UVA altered the expression of phenolic compounds, but our experimental saplings were unlikely to experience photoinhibition because they were not exposed to high levels of light and low temperatures, nor were they nitrogen-limited. We expect that changes in phenolic compounds would have been more pronounced if we had induced photoinhibition.

C₄ photosynthesis allows highly efficient carbon fixation that originates from tightly regulated anatomical and biochemical modifications of leaf architecture. Recent studies showed that leaf transcriptome modifications during leaf ontogeny of closely related C₃ (Tarenaya hassleriana) and C₄ (Gynandropsis gynandra) species within the Cleomaceae family existed but they did not identify any dedicated transcriptional networks or factors specifically driving C₄ leaf ontogeny. RNAseq analysis provides a steady-state quantification of whole-cell mRNAs but does not allow any discrimination between transcriptional and post–transcriptional processes that may occur simultaneously during leaf ontogeny. Here we use exon–intron split analysis (EISA) to determine the extent to which transcriptional and post-transcriptional processes are involved in the regulation of gene expression between young and expanded leaves in both species. C₄-specific changes in post-transcriptional regulation were observed for genes involved in the Calvin–Benson cycle and some photosystem components but not for C₄ core-cycle genes. Overall, this study provides an unbiased genome-wide insight into the post-transcriptional mechanisms that regulate gene expression through the control of mRNA levels and could be central to the onset of C₄ photosynthesis. This mechanism is cytosolic which implies cell-specific modifications of mRNA stability. Understanding this mechanism may be crucial when aiming to transform C₃ crops into C₄ crops.

The seasonality of vegetation, i.e., developmental stages and phenological processes, affects the emission of volatile organic compounds (VOCs). Despite the potential significance, the contributions of seasonality to VOC emission quality and quantity are not well understood and are therefore often ignored in emission simulations. We investigated the VOC emission patterns of young and mature leaves of several Mediterranean plant species in relation to their physiological and developmental changes during the growing period and estimated Es. Foliar emissions of isoprenoids and oxygenated VOCs like methanol and acetone were measured online by means of a proton transfer reaction mass spectrometer (PTR‐MS) and offline with gas chromatography coupled with a mass spectrometer and flame ionization detector. The results suggest that VOC emission is a developmentally regulated process and that quantitative and qualitative variability is plant species specific. Leaf ontogeny clearly influenced both the VOC Es and the relative importance of different VOCs. Methanol was the major compound contributing to the sum of target VOC emissions in young leaves (11.8 ± 10.4 μg g−1 h−1), while its contribution was minor in mature leaves (4.1 ± 4.1 μg g−1 h−1). Several plant species showed a decrease or complete subsidence of monoterpene, sesquiterpene, and acetone emissions upon maturity, perhaps indicating a potential response to the higher defense demands of young emerging leaves. Key Points Developmental stage and phenological processes affect VOC emission Methanol emissions from young leaves were higher than from mature leaves Plants showed decrease or subsidence of monoterpene, sesquiterpene upon maturity

Research highlights: We present evidence indicating that covariation of functional traits among populations of a forest tree is not due to genetic constraints, but rather selective covariance arising from local adaptation to different facets of the climate, namely rainfall and temperature. Background and Aims: Traits frequently covary among natural populations. Such covariation can be caused by pleiotropy and/or linkage disequilibrium, but also may arise when the traits are genetically independent as a direct consequence of natural selection, drift, mutation and/or gene flow. Of particular interest are cases of selective covariance, where natural selection directly generates among-population covariance in a set of genetically independent traits. We here studied the causes of population-level covariation in two key traits in the Australian tree Eucalyptus pauciflora. Materials and Methods: We studied covariation in seedling lignotuber size and vegetative juvenility using 37 populations sampled from throughout the geographic and ecological ranges of E. pauciflora on the island of Tasmania. We integrated evidence from multiple sources: (i) comparison of patterns of trait covariation within and among populations; (ii) climate-trait modelling using machine-learning algorithms; and (iii) selection analysis linking trait variation to field growth in an arid environment. Results: We showed strong covariation among populations compared with the weak genetic correlation within populations for the focal traits. Population differentiation in these genetically independent traits was correlated with different home-site climate variables (lignotuber size with temperature; vegetative juvenility with rainfall), which spatially covaried. The role of selection in shaping the population differentiation in lignotuber size was supported by its relationship with fitness measured in the field. Conclusions: Our study highlights the multi-trait nature of adaptation likely to occur as tree species respond to spatial and temporal changes in climate.