The memory of past water abundance shapes trees 7 years later.

Premise: Tree structure and function are constrained by and acclimate to climatic conditions. Drought limits plant growth and carbon acquisition and can result in "legacy" effects that last beyond the period of water stress. Leaf and twig-level legacy effects of past water abundance, such as that experienced by trees that established under wetter conditions are unknown.

Linking structure to function: the connection between mesophyll structure and intrinsic water use efficiency.

Climate change-driven drought events are becoming unescapable in an increasing number of areas worldwide. Understanding how plants are able to adapt to these changing environmental conditions is a non-trivial challenge. Physiologically, improving a plant's intrinsic water use efficiency (WUEi ) will be essential for plant survival in dry conditions.

Into the Spongy-Verse: Structural Differences between Leaf and Flower Mesophyll.

As the site of almost all terrestrial carbon fixation, the mesophyll tissue is critical to leaf function. However, mesophyll tissue is not restricted only to leaves but also occurs in the laminar, heterotrophic organs of the floral perianth, providing a powerful test of how metabolic differences are linked to differences in tissue structure. Here, we compared mesophyll tissues of leaves and flower perianths of six species using high-resolution X-ray computed microtomography (microCT) imaging.

Analyzing anatomy over three dimensions unpacks the differences in mesophyll diffusive area between sun and shade leaves.

Leaves grown at different light intensities exhibit considerable differences in physiology, morphology and anatomy. Because plant leaves develop over three dimensions, analyses of the leaf structure should account for differences in lengths, surfaces, as well as volumes. In this manuscript, we set out to disentangle the mesophyll surface area available for diffusion per leaf area ( ) into underlying one-, two- and three-dimensional components.

Localized growth and remodelling drives spongy mesophyll morphogenesis.

The spongy mesophyll is a complex, porous tissue found in plant leaves that enables carbon capture and provides mechanical stability. Unlike many other biological tissues, which remain confluent throughout development, the spongy mesophyll must develop from an initially confluent tissue into a tortuous network of cells with a large proportion of intercellular airspace. How the airspace in the spongy mesophyll develops while the tissue remains mechanically stable is unknown.

Desiccation of the leaf mesophyll and its implications for CO diffusion and light processing.

Leaves balance CO and radiative absorption while maintaining water transport to maximise photosynthesis. Related species with contrasting leaf anatomy can provide insights into inherent and stress-induced links between structure and function for commonly measured leaf traits for important crops. We used two walnut species with contrasting mesophyll anatomy to evaluate these integrated exchange processes under non-stressed and drought conditions using a combination of light microscopy, X-ray microCT, gas exchange, hydraulic conductance, and chlorophyll distribution profiles through leaves.

The three-dimensional construction of leaves is coordinated with water use efficiency in conifers.

Conifers prevail in the canopies of many terrestrial biomes, holding a great ecological and economic importance globally. Current increases in temperature and aridity are imposing high transpirational demands and resulting in conifer mortality. Therefore, identifying leaf structural determinants of water use efficiency is essential for predicting physiological impacts due to environmental variation.

Understanding airspace in leaves: 3D anatomy and directional tortuosity.

The leaf intercellular airspace is a tortuous environment consisting of cells of different shapes, packing densities, and orientation, all of which have an effect on the travelling distance of molecules from the stomata to the mesophyll cell surfaces. Tortuosity, the increase in displacement over the actual distance between two points, is typically defined as encompassing the whole leaf airspace, but heterogeneity in pore dimensions and orientation between the spongy and palisade mesophyll likely result in heterogeneity in tortuosity along different axes and would predict longer traveling distance along the path of least tortuosity, such as vertically within the columnar cell matrix of the palisade layer. Here, we compare a previously established geometric method to a random walk approach, novel for this analysis in plant leaves, in four different Eucalyptus species.

A meta-analysis of mesophyll conductance to CO2 in relation to major abiotic stresses in poplar species.

Mesophyll conductance (gm) determines the diffusion of CO2 from the substomatal cavities to the site of carboxylation in the chloroplasts and represents a critical component of the diffusive limitation of photosynthesis. In this study, we evaluated the average effect sizes of different environmental constraints on gm in Populus spp., a forest tree model.

Maximum CO diffusion inside leaves is limited by the scaling of cell size and genome size.

Maintaining high rates of photosynthesis in leaves requires efficient movement of CO from the atmosphere to the mesophyll cells inside the leaf where CO is converted into sugar. CO diffusion inside the leaf depends directly on the structure of the mesophyll cells and their surrounding airspace, which have been difficult to characterize because of their inherently three-dimensional organization. Yet faster CO diffusion inside the leaf was probably critical in elevating rates of photosynthesis that occurred among angiosperm lineages.

Assessing adaptive and plastic responses in growth and functional traits in a 10-year-old common garden experiment with pedunculate oak ( L.) suggests that directional selection can drive climatic adaptation.

Understanding how tree species will respond to a future climate requires reliable and quantitative estimates of intra-specific variation under current climate conditions. We studied three 10-year-old common garden experiments established across a rainfall and drought gradient planted with nearly 10,000 pedunculate oak ( L.) trees from ten provenances with known family structure.

Digitally deconstructing leaves in 3D using X-ray microcomputed tomography and machine learning.

Premise: X-ray microcomputed tomography (microCT) can be used to measure 3D leaf internal anatomy, providing a holistic view of tissue organization. Previously, the substantial time needed for segmenting multiple tissues limited this technique to small data sets, restricting its utility for phenotyping experiments and limiting our confidence in the inferences of these studies due to low replication numbers.

Methods And Results: We present a Python codebase for random forest machine learning segmentation and 3D leaf anatomical trait quantification that dramatically reduces the time required to process single-leaf microCT scans into detailed segmentations.

Shape matters: the pitfalls of analyzing mesophyll anatomy.

This article is a Commentary on https://doi.org/10.1111/nph.

Beyond Porosity: 3D Leaf Intercellular Airspace Traits That Impact Mesophyll Conductance.

The leaf intercellular airspace (IAS) is generally considered to have high conductance to CO diffusion relative to the liquid phase. While previous studies accounted for leaf-level variation in porosity and mesophyll thickness, they omitted 3D IAS traits that potentially influence IAS conductance (g). Here, we reevaluated the standard equation for g by incorporating tortuosity, lateral path lengthening, and IAS connectivity.

The bias of a two-dimensional view: comparing two-dimensional and three-dimensional mesophyll surface area estimates using noninvasive imaging.

The mesophyll surface area exposed to intercellular air space per leaf area (S ) is closely associated with CO diffusion and photosynthetic rates. S is typically estimated from two-dimensional (2D) leaf sections and corrected for the three-dimensional (3D) geometry of mesophyll cells, leading to potential differences between the estimated and actual cell surface area. Here, we examined how 2D methods used for estimating S compare with 3D values obtained from high-resolution X-ray microcomputed tomography (microCT) for 23 plant species, with broad phylogenetic and anatomical coverage.

Temperature gradients assist carbohydrate allocation within trees.

Trees experience two distinct environments: thermally-variable air and thermally-buffered soil. This generates intra-tree temperature gradients, which can affect carbon metabolism and water transport. In this study, we investigated whether carbohydrate allocation within trees is assisted by temperature gradients.

Excess Diffuse Light Absorption in Upper Mesophyll Limits CO Drawdown and Depresses Photosynthesis.

In agricultural and natural systems, diffuse light can enhance plant primary productivity due to deeper penetration into and greater irradiance of the entire canopy. However, for individual sun-grown leaves from three species, photosynthesis is actually less efficient under diffuse compared with direct light. Despite its potential impact on canopy-level productivity, the mechanism for this leaf-level diffuse light photosynthetic depression effect is unknown.

The light response of mesophyll conductance is controlled by structure across leaf profiles.

Mesophyll conductance to CO (g ) may respond to light either through regulated dynamic mechanisms or due to anatomical and structural factors. At low light, some layers of cells in the leaf cross-section approach photocompensation and contribute minimally to bulk leaf photosynthesis and little to whole leaf g (g ). Thus, the bulk g will appear to respond to light despite being based upon cells having an anatomically fixed mesophyll conductance.

Greater efficiency of water use in poplar clones having a delayed response of mesophyll conductance to drought.

Improvement of water use efficiency is a key objective to improve the sustainability of cultivated plants, especially fast growing species with high water consumption like poplar. It is well known that water use efficiency (WUE) varies considerably among poplar genotypes, and it was recently suggested that the use of the mesophyll-to-stomatal conductance ratio (gm/gs) would be an appropriate trait to improve WUE. The responses of 7-week-old cuttings of four hybrid poplar clones and one native Balsam poplar (Populus balsamifera L.

Threshold response of mesophyll CO2 conductance to leaf hydraulics in highly transpiring hybrid poplar clones exposed to soil drying.

Mesophyll conductance (gm) has been shown to impose significant limitations to net CO2 assimilation (A) in various species during water stress. Net CO2 assimilation is also limited by stomatal conductance to water (gsw), both having been shown to co-vary with leaf hydraulic conductance (Kleaf). Lately, several studies have suggested a close functional link between Kleaf, gsw, and gm.