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In plants, the delivery of the products of photosynthesis is achieved through a hydraulic system labeled as phloem. This semi-permeable plant tissue consists of living cells that contract and expand in response to fluid pressure and flow velocity fluctuations. The Münch pressure flow theory, which is based on osmosis providing the necessary pressure gradient to drive the mass flow of carbohydrates, is currently the most accepted model for such sucrose transport. When this hypothesis is combined with the conservation of fluid mass and momentum as well as sucrose mass, many simplifications must be invoked to mathematically close the problem and to resolve the flow. This study revisits such osmotically driven flows by developing a new two-dimensional numerical model in cylindrical coordinates for an elastic membrane and a concentration-dependent viscosity. It is demonstrated that the interaction between the hydrodynamic and externally supplied geometrical characteristic of the phloem has a significant effect on the front speed of sucrose transport. These results offer a novel perspective about the evolutionary adaptation of plant hydraulic traits to optimize phloem soluble compounds transport efficiency. 

Key Points The physics of sugar translocation in phloem cells is complicated by their elasticity Membrane elasticity leads to a slower front speed when compared to a theoretical rigid membrane The role of sieve plates, porous media that exist within the phloem, is reconsidered to be structural instead of hydraulic 

ABSTRACT AimTo quantify the intra‐community variability of leaf‐out (ICVLo) among dominant trees in temperate deciduous forests, assess its links with specific and phylogenetic diversity, identify its environmental drivers and deduce its ecological consequences with regard to radiation received and exposure to late frost. LocationEastern North America (ENA) and Europe (EUR). Time Period2009–2022. Major Taxa StudiedTemperate deciduous forest trees. MethodsWe developed an approach to quantify ICVLo through the analysis of RGB images taken from phenological cameras. We related ICVLo to species richness, phylogenetic diversity and environmental conditions. We quantified the intra‐community variability of the amount of radiation received and of exposure to late frost. ResultsLeaf‐out occurred over a longer time interval in ENA than in EUR. The sensitivity of leaf‐out to temperature was identical in both regions (−3.4 days per °C). The distributions of ICVLo were similar in EUR and ENA forests, despite the latter being more species‐rich and phylogenetically diverse. In both regions, cooler conditions and an earlier occurrence of leaf‐out resulted in higher ICVLo. ICVLo resulted in ca. 8% difference of radiation received from leaf‐out to September among individual trees. Forest communities in ENA had shorter safety margins as regards the exposure to late frosts, and were actually more frequently exposed to late frosts. Main ConclusionsWe conducted the first intercontinental analysis of the variability of leaf‐out at the scale of tree communities. North American and European forests showed similar ICVLo, in spite of their differences in terms of species richness and phylogenetic diversity, highlighting the relevance of environmental controls on ICVLo. We quantified two ecological implications of ICVLo (difference in terms of radiation received and exposure to late frost), which should be explored in the context of ongoing climate change, which affects trees differently according to their phenological niche. 

Abstract 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. 

1. Wood anatomical traits shape a xylem segment's hydraulic efficiency and embolism spread resistance due to declining water potential. It has been known for decades that variations in conduit connectivity play a role in altering xylem hydraulics. However, evaluating the precise effect of conduit connectivity has been elusive. The objective is to establish an analytical linkage between conduit connectivity and grouping and tissue-scale hydraulics. 2. It is hypothesized that an increase in conduit connectivity brings improved resistance to embolism spread due to increased hydraulic pathway redundancy. However, an increase in conduit connectivity could also reduce resistance due to increased embolism spread speed with respect to pressure. We elaborate on this trade-off using graph theory, percolation theory, and computational modeling of xylem. The results are validated using anatomical measurements of Acer branch xylem. 3. Considering only species with vessels, increases in connectivity improve resistance to embolism spread without negatively affecting hydraulic conductivity. The often measured grouping index fails to capture the totality of the effect of conduit connectivity on xylem hydraulics. 4. Variations in xylem network characteristics, such as conduit connectivity, might explain why hypothesized trends among woody species, like the ‘safety-efficiency’ trade-off hypothesis, are weaker than expected.