Transcriptome and physiological analyses provide insights into the leaf epicuticular wax accumulation mechanism in yellowhorn

Plantations and production of yellowhorn, one of the most important woody oil and urban greening trees widely cultivated in northern China, have gradually become limited by drought stress. The epicuticular wax layer plays a key role in the protection of yellowhorn trees from drought and other stresses. However, there is no research on the mechanism of wax loading in yellowhorn trees. In this study, we investigated the anatomical and physiological characteristics of leaves from different germplasm resources and different parts of the same tree and compared their cuticle properties. In addition, the different expression patterns of genes involved in wax accumulation were analyzed, and a coexpression network was built based on transcriptome sequencing data. Morphological and physiological comparisons found that the sun leaves from the outer part of the crown had thicker epicuticular wax, which altered the permeability and improved the drought resistance of leaves, than did shade leaves. Based on transcriptome data, a total of 3008 and 1324 differentially expressed genes (DEGs) were identified between the sun leaves and shade leaves in glossy- and non-glossy-type germplasm resources, respectively. We identified 138 DEGs involved in wax biosynthesis and transport, including structural genes (such as LACS8, ECH1, and ns-LTP) and transcription factors (such as MYB, WRKY, and bHLH transcription factor family proteins). The coexpression network showed a strong correlation between these DEGs. The differences in gene expression patterns between G- and NG-type germplasm resources under different light conditions were very clear. These results not only provide a theoretical basis for screening and developing drought-resistant yellowhorn germplasm resources but also provide a data platform to reveal the wax accumulation process of yellowhorn leaves.

Can improved canopy light transmission ameliorate loss of photosynthetic efficiency in the shade? An investigation of natural variation in Sorghum bicolor.

Previous studies have found that maximum quantum yield of CO2 assimilation (ΦCO2,max,app) declines in lower canopies of maize and Miscanthus, a maladaptive response to self-shading. These observations were limited to single genotypes, leaving it unclear that the maladaptive shade response is a general property of this C4 grass tribe, the Andropogoneae. We explored the generality of this maladaptation by testing the hypothesis that erect leaf forms (erectophiles), which allow more light into the lower canopy, suffer less of a decline in photosynthetic efficiency than drooping leaf (planophile) forms. On average, ΦCO2,max,app declined 27% in lower canopy leaves across 35 accessions, but the decline was over twice as great in planophiles than in erectophiles. The loss of photosynthetic efficiency involved a decoupling between electron transport and assimilation. This was not associated with increased bundle sheath leakage, based on 13C measurements. In both planophiles and erectophiles, shaded leaves had greater leaf absorptivity and lower activities of key C4 enzymes than sun leaves. The erectophile form is considered more productive because it allows a more effective distribution of light through the canopy to support photosynthesis. We show that in sorghum, it provides a second benefit, maintenance of higher ΦCO2,max,app to support efficient use of that light resource.

Making Decisions with Data

Reliable scientific conclusions are based on verifiable empirical evidence. But data must be transformed and interpreted before they become evidence, and statistical inference plays an important role in the process of interpretation. Biologists use statistics to organize and analyze data so that they can make inferences and use the data as evidence. Students should have opportunities to collect and analyze data in their biology classes as well. In this activity, students collect data on the surface areas of sun leaves and shade leaves, then analyze the data using the independentsamples t-test. The t-test procedure can be used in investigations where two groups are compared on one dependent variable.

Differences in Characteristics of Photosynthesis and Nitrogen Utilization in Leaves of the Black Locust (Robinia pseudoacacia L.) According to Leaf Position

Robinia pseudoacacia L. has been widely planted worldwide for a variety of purposes, but it is a nonindigenous species currently invading the central part of Japanese river terraces. To understand and control this invasion, we investigated how this species invests nitrogen resources in different functions depending on the leaf location, and how these resources are used in physiological reactions such as photosynthesis. The Tama river terrace was examined in Tokyo, Japan. The leaf nitrogen (N) concentration, chlorophyll (Chl) concentration, Chl a/b ratio, leaf mass per unit area (LMA) and ribulose-1,5-bisphosphate carboxylase oxygenase (RuBisCo) concentration were all significantly lower in shade leaves than in leaves exposed to the sun. Conversely, the net photosynthetic rate in saturated light conditions (Pmax), the net photosynthetic rate under enhanced CO2 concentration and light saturation (Amax), the maximum carboxylation rate of RuBisCo (Vcmax) and the maximum rate of electron transport driving RUBP regeneration (Jmax) were all significantly lower in shade leaves than in leaves exposed to the sun. We also found that RuBisCo/N and Chl/N were significantly less in shade leaves, and values of Jmax/N, Vcmax/N less in shade leaves than in sun leaves, but not significantly. Allocation of nitrogen in leaves to photosynthetic proteins, RuBisCo (NR) was broadly less in shade leaves, and NL (light-harvesting complex: LHC, photosystem I and II: PSI and PSII) and NE (electron transport) were also lower. The N remaining was much greater in shade leaves than in sun leaves. We suggest that N remobilization from RuBisCo is more efficient than remobilization from proteins of NE, and from NL. This study shows that R. pseudoacacia has an enhanced ability to adapt to environmental changes via characteristic changes in N allocation trade-offs and physiological traits in its sun and shade leaves.

Foliar sampling with an unmanned aerial system (UAS) reveals spectral and functional trait differences within tree crowns

Imaging spectroscopy is currently the best approach for continuously mapping forest canopy traits, which is important for ecosystem and biodiversity assessments. Ideally, models are trained with trait data from fully sunlit leaves from the top of the canopy. However, sampling leaves at the top of the canopy is often difficult, and sunlit foliage from the crown periphery is collected instead, assuming minimal within-crown trait variation among sunlit leaves. We tested the degree to which crown position affects foliar traits and spectra using mixed-effects models comparing sun leaves from the crown centre of mature sugar maple (Acer saccharum Marsh.) trees collected with DeLeaves, a novel twig-sampling unmanned aerial system device, with sun leaves from the crown periphery collected with a pole pruner. Sun leaves from the crown centre differed from sun leaves from the crown periphery in absorption, reflectance, transmittance, and a series of foliar traits, including leaf thickness, leaf mass, and leaf nitrogen content per unit area, demonstrating differences in resource allocation depending on sun exposure. Our study highlights the importance of accurately matching the location of foliar samples and spectral data and of sampling across gradients of intra-individual variation for the accurate prediction of foliar trait distributions across and within canopies using imaging spectroscopy.

Light regimen-induced variability of photosynthetic pigments and UV-B absorbing compounds in Luzula sylvatica from Arcto-Alpine tundra

The aim of this study was to evaluate the effects of different in situ light regimen on ecophysiological parameters of Luzula sylvatica leaves. Plants of L. sylvatica grown under natural sunny and shade conditions in arcto-alpine tundra were analyzed with respect to their leaf anatomy, content of photosynthetic pigments, UV absorbing compounds and phenanthrenoid compounds. Relationship between chlorophyll concentrations (Chla+b) and SPAD values was determined for sun and shade leaves measured repeatedly within summer and autumn seasons 2019 and 2020. Pooled data showed curvilinear Chla+b to SPAD relationship with the highest Chla+b and SPAD values found for shade leaves. Sun leaves had higher UV-B absorbing compounds contents than shade ones. The HPLC-DAD analysis revealed significant amount of soluble flavonoids in Luzula sylvatica leaves, amongst others the flavone-luteolin and its derivatives (e.g. tentatively identified luteolin-methyl-glucoside and luteolin-glucoside). The accumulation of luteolin based compounds in sun acclimated leaves is also plausible explanation for the higher antioxidant activity determined in sun leaf extraxts. Such response of flavonoid metabolism may help L.S. to cope with excessive-light stress through UV-attenuation mechanism and ROS scavanging. Additionally, phenanthrenoid compounds contents in L. sylvatica leaves were determined. Altogether, 9 phenanthrenoid compounds were identified by HPLC-HRMS. Their content was markedly different (up to the factor of 5) between sun and shade leaves of L.sylvatica.

Similar temperature dependence of photosynthetic parameters in sun and shade leaves of three tropical tree species

Photosynthetic carbon uptake by tropical forests is of critical importance in regulating the earth’s climate, but rising temperatures threaten this stabilizing influence of tropical forests. Most research on how temperature affects photosynthesis focuses on fully sun-exposed leaves, and little is known about shade leaves, even though shade leaves greatly outnumber sun leaves in lowland tropical forests. We measured temperature responses of light-saturated photosynthesis, stomatal conductance, and the biochemical parameters VCMax (maximum rate of RuBP carboxylation) and JMax (maximum rate of RuBP regeneration, or electron transport) on sun and shade leaves of mature tropical trees of three species in Panama. As expected, biochemical capacities and stomatal conductance were much lower in shade than in sun leaves, leading to lower net photosynthesis rates. However, the key temperature response traits of these parameters—the optimum temperature (TOpt) and the activation energy—did not differ systematically between sun and shade leaves. Consistency in the JMax to VCMax ratio further suggested that shade leaves are not acclimated to lower temperatures. For both sun and shade leaves, stomatal conductance had the lowest temperature optimum (~25 °C), followed by net photosynthesis (~30 °C), JMax (~34 °C) and VCMax (~38 °C). Stomatal conductance of sun leaves decreased more strongly with increasing vapor pressure deficit than that of shade leaves. Consistent with this, modeled stomatal limitation of photosynthesis increased with increasing temperature in sun but not shade leaves. Collectively, these results suggest that modeling photosynthetic carbon uptake in multi-layered canopies does not require independent parameterization of the temperature responses of the biochemical controls over photosynthesis of sun and shade leaves. Nonetheless, to improve the representation of the shade fraction of carbon uptake dynamics in tropical forests, better understanding of stomatal sensitivity of shade leaves to temperature and vapor pressure deficit will be required.

Biomimetic Groundwork for Thermal Exchange Structures Inspired by Plant Leaf Design

Geometry is a determining factor for thermal performance in both biological and technical systems. While biology has inspired thermal design before, biomimetic translation of leaf morphology into structural aspects of heat exchangers remains largely unaddressed. One determinant of plant thermal endurance against environmental exposure is leaf shape, which modulates the leaf boundary layer, transpiration, evaporative cooling, and convective exchange. Here, we lay the research groundwork for the extraction of design principles from leaf shape relations to heat and mass transfer. Leaf role models were identified from an extensive literature review on environmentally sensitive morphology patterns and shape-dependent exchange. Addressing canopy sun–shade dimorphism, sun leaves collected from multiple oak species exceeded significantly in margin extension and shape dissection. Abstracted geometries (i.e., elongated; with finely toothed edges; with few large-scale teeth) were explored with paper models of the same surface area in a controlled environment of minimal airflow, which is more likely to induce leaf thermal stress. For two model characteristic dimensions, evaporation rates were significantly faster for the dissected geometries. Shape-driven transfer enhancements were higher for the smaller models, and finely toothed edges reached local cooling up to 10 °C below air temperature. This investigation breaks new ground for solution-based biomimetics to inform the design of evaporation-assisted and passively enhanced thermal systems.

Incident Light and Leaf Age Influence Leaflet Element Concentrations of Cycas micronesica Trees

The need for improved knowledge on conservation and management of cycad species has generated recent interest in compiling a database on leaf nutrient concentrations. However, the sampling protocols have not been consistent among reports and the influences of some plant and habitat traits on the plasticity of cycad leaf nutrient concentrations has not been adequately determined. We used Cycas micronesica K.D. Hill trees to determine the role of incident light level and leaf age on leaflet content of 11 essential elements. Shade leaves exhibited increased mass-based concentration for nitrogen, phosphorus, and potassium above that of sun leaves. Shade leaves exhibited decreased area-based concentration for all of the macro- and micronutrients below that of sun leaves. Mass-based concentration of nitrogen, phosphorus, and potassium decreased with leaf age, and that of calcium, magnesium, iron, manganese, and zinc increased with leaf age. These findings indicate the relative leaf age and the amount of shade or incident light at the leaf level must be recorded and reported for leaf tissue studies in cycads in order to reduce ambiguity and ensure repeatability.