Freeze/Thaw-Induced Embolism: Probability of Critical Bubble Formation Depends on Speed of Ice Formation

Bubble formation in the conduits of woody plants sets a challenge for uninterrupted water transportation from the soil up to the canopy. Freezing and thawing of stems has been shown to increase the number of air-filled (embolized) conduits, especially in trees with large conduit diameters. Despite numerous experimental studies, the mechanisms leading to bubble formation during freezing have not been addressed theoretically. We used classical nucleation theory and fluid mechanics to show which mechanisms are most likely to be responsible for bubble formation during freezing and what parameters determine the likelihood of the process. Our results confirm the common assumption that bubble formation during freezing is most likely due to gas segregation by ice. If xylem conduit walls are not permeable to the salts expelled by ice during the freezing process, osmotic pressures high enough for air seeding could be created. The build-up rate of segregated solutes in front of the ice-water interface depends equally on conduit diameter and freezing velocity. Therefore, bubble formation probability depends on these variables. The dependence of bubble formation probability on freezing velocity means that the experimental results obtained for cavitation threshold conduit diameters during freeze/thaw cycles depend on the experimental setup; namely sample size and cooling rate. The velocity dependence also suggests that to avoid bubble formation during freezing trees should have narrow conduits where freezing is likely to be fast (e.g., branches or outermost layer of the xylem). Avoidance of bubble formation during freezing could thus be one piece of the explanation why xylem conduit size of temperate and boreal zone trees varies quite systematically.We are grateful for support from the Andrew W. Mellon Foundation and the Materials Research Science and Engineering Center at Harvard University. We also thank the Australian Research Council for support (DP110105380)

Ion Induced Changes in the Structure of Bordered Pit Membranes

Ion-mediated changes in xylem hydraulic resistance are hypothesized to result from hydrogel like properties of pectins located in the bordered pit membranes separating adjacent xylem vessels. Although the kinetics of the ion-mediated changes in hydraulic resistance are consistent with the swelling/deswelling behavior of pectins, there is no direct evidence of this activity. In this report we use atomic force microscopy (AFM) to investigate structural changes in bordered pit membranes associated with changes in the ionic concentration of the surrounding solution. When submerged in de-ionized water, AFM revealed bordered pit membranes as relatively smooth, soft, and lacking any sharp edges surface, in contrast to pictures from scanning electron microscope (SEM) or AFM performed on air-dry material. Exposure of the bordered pit membranes to 50 mM KCl solution resulted in significant changes in both surface physical properties and elevation features. Specifically, bordered pit membranes became harder and the fiber edges were clearly visible. In addition, the membrane contracted and appeared much rougher due to exposed microfibers. In neither solution was there any evidence of discrete pores through the membrane whose dimensions were altered in response to the ionic composition of the surrounding solution. Instead the variable hydraulic resistance appears to involve changes in the both the permeability and the thickness of the pit membrane

Optimal concentrations in transport systems

Many biological and man-made systems rely on transport systems for the distribution of material, for example matter and energy. Material transfer in these systems is determined by the flow rate and the concentration of material. While the most concentrated solutions offer the greatest potential in terms of material transfer, impedance typically increases with concentration, thus making them the most difficult to transport. We develop a general framework for describing systems for which impedance increases with concentration, and consider material flow in four different natural systems: blood flow in vertebrates, sugar transport in vascular plants and two modes of nectar drinking in birds and insects. The model provides a simple method for determining the optimum concentration c[subscript opt] in these systems. The model further suggests that the impedance at the optimum concentration μ[subscript opt] may be expressed in terms of the impedance of the pure (c = 0) carrier medium μ[subscript 0] as μ[subscript opt]∼2[superscript α]μ[subscript 0], where the power α is prescribed by the specific flow constraints, for example constant pressure for blood flow (α = 1) or constant work rate for certain nectar-drinking insects (α = 6). Comparing the model predictions with experimental data from more than 100 animal and plant species, we find that the simple model rationalizes the observed concentrations and impedances. The model provides a universal framework for studying flows impeded by concentration, and yields insight into optimization in engineered systems, such as traffic flow.National Science Foundation (U.S.) (Grant 1021779)National Science Foundation (U.S.) (Grant DMS-0907955)National Science Foundation (U.S.). Materials Research Science and Engineering Centers (Program) (Grant DMR-0820484

Leaf Carbon Export and Nonstructural Carbohydrates in Relation to Diurnal Water Dynamics in Mature Oak Trees

Trees typically experience large diurnal depressions in water potential, which may impede carbon export from leaves during the day because the xylem is the source of water for the phloem. As water potential becomes more negative, higher phloem osmotic concentrations are needed to draw water in from the xylem. Generating this high concentration of sugar in the phloem is particularly an issue for the ∼50% of trees that exhibit passive loading. These ideas motivate the hypothesis that carbon export in woody plants occurs predominantly at night, with sugars that accumulate during the day assisting in mesophyll turgor maintenance or being converted to starch. To test this, diurnal and seasonal patterns of leaf nonstructural carbohydrates, photosynthesis, solute, and water potential were measured, and carbon export was estimated in leaves of five mature (\u3e20 m tall) red oak (Quercus rubra) trees, a species characterized as a passive loader. Export occurred throughout the day at equal or higher rates than at night despite a decrease in water potential to −1.8 MPa at midday. Suc and starch accumulated over the course of the day, with Suc contributing ∼50% of the 0.4 MPa diurnal osmotic adjustment. As a result of this diurnal osmotic adjustment, estimates of midday turgor were always \u3e0.7 MPa. These findings illustrate the robustness of phloem functioning despite diurnal fluctuations in leaf water potential and the role of nonstructural carbohydrates in leaf turgor maintenance

Modelled hydraulic redistribution by sunflower (Helianthus annuus L.) matches observed data only after including night-time transpiration

Author Posting. © The Author(s), 2013. This is the author's version of the work. It is posted here by permission of John Wiley & Sons for personal use, not for redistribution. The definitive version was published in Plant, Cell & Environment 37 (2014): 899-910, doi:10.1111/pce.12206.The movement of water from moist to dry soil layers through the root systems of plants, referred to as hydraulic redistribution (HR), occurs throughout the world and is thought to influence carbon and water budgets and ecosystem functioning. The realized hydrologic, biogeochemical, and ecological consequences of HR depend on the amount of redistributed water, while the ability to assess these impacts requires models that correctly capture HR magnitude and timing. Using several soil types and two eco-types of sunflower (Helianthus annuus L.) in split-pot experiments, we examined how well the widely used HR modeling formulation developed by Ryel et al. (2002) matched experimental determination of HR across a range of water potential driving gradients. H. annuus carries out extensive nighttime transpiration, and though over the last decade it has become more widely recognized that nighttime transpiration occurs in multiple species and many ecosystems, the original Ryel et al. (2002) formulation does not include the effect of nighttime transpiration on HR. We developed and added a representation of nighttime transpiration into the formulation, and only then was the model able to capture the dynamics and magnitude of HR we observed as soils dried and nighttime stomatal behavior changed, both influencing HR.This work was supported by a NOAA Climate and Global Change Postdoctoral Fellowship to RBN, administered by the University Corporation for Atmospheric Research, by a grant from the Andrew W. Mellon Foundation to NMH, and by DOE Terrestrial Ecosystem Science grant ER65389 to ZGC and RBN.2014-10-2

Relationship between Hexokinase and the Aquaporin PIP1 in the Regulation of Photosynthesis and Plant Growth

Increased expression of the aquaporin NtAQP1, which is known to function as a plasmalemma channel for CO2 and water, increases the rate of both photosynthesis and transpiration. In contrast, increased expression of Arabidopsis hexokinase1 (AtHXK1), a dual-function enzyme that mediates sugar sensing, decreases the expression of photosynthetic genes and the rate of transpiration and inhibits growth. Here, we show that AtHXK1 also decreases root and stem hydraulic conductivity and leaf mesophyll CO2 conductance (gm). Due to their opposite effects on plant development and physiology, we examined the relationship between NtAQP1 and AtHXK1 at the whole-plant level using transgenic tomato plants expressing both genes simultaneously. NtAQP1 significantly improved growth and increased the transpiration rates of AtHXK1-expressing plants. Reciprocal grafting experiments indicated that this complementation occurs when both genes are expressed simultaneously in the shoot. Yet, NtAQP1 had only a marginal effect on the hydraulic conductivity of the double-transgenic plants, suggesting that the complementary effect of NtAQP1 is unrelated to shoot water transport. Rather, NtAQP1 significantly increased leaf mesophyll CO2 conductance and enhanced the rate of photosynthesis, suggesting that NtAQP1 facilitated the growth of the double-transgenic plants by enhancing mesophyll conductance of CO2

Impacts of elevated atmospheric CO₂ on nutrient content of important food crops.

One of the many ways that climate change may affect human health is by altering the nutrient content of food crops. However, previous attempts to study the effects of increased atmospheric CO2 on crop nutrition have been limited by small sample sizes and/or artificial growing conditions. Here we present data from a meta-analysis of the nutritional contents of the edible portions of 41 cultivars of six major crop species grown using free-air CO2 enrichment (FACE) technology to expose crops to ambient and elevated CO2 concentrations in otherwise normal field cultivation conditions. This data, collected across three continents, represents over ten times more data on the nutrient content of crops grown in FACE experiments than was previously available. We expect it to be deeply useful to future studies, such as efforts to understand the impacts of elevated atmospheric CO2 on crop macro- and micronutrient concentrations, or attempts to alleviate harmful effects of these changes for the billions of people who depend on these crops for essential nutrients

On measuring the response of mesophyll conductance to carbon dioxide with the variable J method

The response of mesophyll conductance to CO2 (gm) to environmental variation is a challenging parameter to measure with current methods. The ‘variable J’ technique, used in the majority of studies of gm, assumes a one-to-one relationship between photosystem II (PSII) fluorescence and photosynthesis under non-photorespiratory conditions. When calibrating this relationship for Populus trichocarpa, it was found that calibration relationships produced using variation in light and CO2 were not equivalent, and in all cases the relationships were non-linear—something not accounted for in previous studies. Detailed analyses were performed of whether different calibration procedures affect the observed gm response to CO2. Past linear and assumed calibration methods resulted in systematic biases in the fluorescence estimates of electron transport. A sensitivity analysis on modelled data (where gm was held constant) demonstrated that biases in the estimation of electron transport as small as 2% (∼0.5 μmol m−2 s−1) resulted in apparent changes in the relationship of gm to CO2 of similar shape and magnitude to those observed with past calibration techniques. This sensitivity to biases introduced during calibrations leads to results where gm artefactually decreases with CO2, assuming that gm is constant; if gm responds to CO2, then biases associated with past calibration methods would lead to overestimates of the slope of the relationship. Non-linear calibrations were evaluated; these removed the bias present in past calibrations, but the method remained sensitive to measurement errors. Thus measurement errors, calibration non-linearities leading to bias, and the sensitivity of variable J gm hinders its use under conditions of varying CO2 or light