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Creators/Authors contains: "Rockwell, Fulton E."

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  1. Abstract

    The water status of the living tissue in leaves between the xylem and stomata (outside xylem zone (OXZ) plays a critical role in plant function and global mass and energy balance but has remained largely inaccessible. We resolve the local water relations of OXZ tissue using a nanogel reporter of water potential (ψ), AquaDust, that enables an in situ, nondestructive measurement of both ψ of xylem and highly localized ψ at the terminus of transpiration in the OXZ. Working in maize (Zea mays L.), these localized measurements reveal gradients in the OXZ that are several folds larger than those based on conventional methods and values of ψ in the mesophyll apoplast well below the macroscopic turgor loss potential. We find a strong loss of hydraulic conductance in both the bundle sheath and the mesophyll with decreasing xylem potential but not with evaporative demand. Our measurements suggest the OXZ plays an active role in regulating the transpiration path, and our methods provide the means to study this phenomenon.

     
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  2. Abstract Background

    Recent reports of extreme levels of undersaturation in internal leaf air spaces have called into question one of the foundational assumptions of leaf gas exchange analysis, that leaf air spaces are effectively saturated with water vapour at leaf surface temperature. Historically, inferring the biophysical states controlling assimilation and transpiration from the fluxes directly measured by gas exchange systems has presented a number of challenges, including: (1) a mismatch in scales between the area of flux measurement, the biochemical cellular scale and the meso-scale introduced by the localization of the fluxes to stomatal pores; (2) the inaccessibility of the internal states of CO2 and water vapour required to define conductances; and (3) uncertainties about the pathways these internal fluxes travel. In response, plant physiologists have adopted a set of simplifying assumptions that define phenomenological concepts such as stomatal and mesophyll conductances.

    Scope

    Investigators have long been concerned that a failure of basic assumptions could be distorting our understanding of these phenomenological conductances, and the biophysical states inside leaves. Here we review these assumptions and historical efforts to test them. We then explore whether artefacts in analysis arising from the averaging of fluxes over macroscopic leaf areas could provide alternative explanations for some part, if not all, of reported extreme states of undersaturation.

    Conclusions

    Spatial heterogeneities can, in some cases, create the appearance of undersaturation in the internal air spaces of leaves. Further refinement of experimental approaches will be required to separate undersaturation from the effects of spatial variations in fluxes or conductances. Novel combinations of current and emerging technologies hold promise for meeting this challenge.

     
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  3. Abstract

    A frequently expressed viewpoint across the Earth science community is that global soil moisture estimates from satellite L‐band (1.4 GHz) measurements represent moisture only in a shallow surface layer (0–5 cm) and consequently are of limited value for studying global terrestrial ecosystems because plants use water from deeper rootzones. Using this argumentation, many observation‐based land surface studies avoid satellite‐observed soil moisture. Here, based on peer‐reviewed literature across several fields, we argue that such a viewpoint is overly limiting for two reasons. First, microwave soil emission depth considerations and statistical considerations of vertically correlated soil moisture information together indicate that L‐band measurements carry information about soil moisture extending below the commonly referenced 5 cm in many conditions. However, spatial variations of effective depths of representation remain uncertain. Second, in reviewing isotopic tracer field studies of plant water uptake, we find a prevalence of vegetation that primarily draws moisture from these upper soil layers. This is especially true for grasslands and croplands covering more than a third of global vegetated surfaces. Even some deeper‐rooted species (i.e., shrubs and trees) preferentially or seasonally draw water from the upper soil layers. Therefore, L‐band satellite soil moisture estimates are more relevant to global vegetation water uptake than commonly appreciated (i.e., relevant beyond only shallow soil processes like soil evaporation). Our commentary encourages the application of satellite soil moisture across a broader range of terrestrial hydrosphere and biosphere studies while urging more rigorous estimates of its effective depth of representation.

     
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  4. Summary

    The hydraulic system of vascular plants and its integrity is essential for plant survival. To transport water under tension, the walls of xylem conduits must approximate rigid pipes. Against this expectation, conduit deformation has been reported in the leaves of a few species and hypothesized to function as a ‘circuit breaker’ against embolism. Experimental evidence is lacking, and its generality is unknown.

    We demonstrated the role of conduit deformation in protecting the upstream xylem from embolism through experiments on three species and surveyed a diverse selection of vascular plants for conduit deformation in leaves.

    Conduit deformation in minor veins occurred before embolism during slow dehydration. When leaves were exposed to transient increases in transpiration, conduit deformation was accompanied by large water potential differences from leaf to stem and minimal embolism in the upstream xylem. In the three species tested, collapsible vein endings provided clear protection of upstream xylem from embolism during transient increases in transpiration.

    We found conduit deformation in diverse vascular plants, including 11 eudicots, ginkgo, a cycad, a fern, a bamboo, and a grass species, but not in two bamboo and a palm species, demonstrating that the potential for ‘circuit breaker’ functionality may be widespread across vascular plants.

     
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  5. Premise

    The dimensions of phloem sieve elements have been shown to vary as a function of tree height, decreasing hydraulic resistance as the transport pathway lengthens. However, little is known about ontogenetic patterns of sieve element scaling. Here we examine within a single species (Quercus rubra) how decreases in hydraulic resistance with distance from the plant apex are mediated by overall plant size.

    Methods

    We sampled and imaged phloem tissue at multiple heights along the main stem and in the live crown of four size classes of trees using fluorescence and scanning electron microscopy. Sieve element length and radius, the number of sieve areas per compound plate, pore number, and pore radius were used to calculate total hydraulic resistance at each sampling location.

    Results

    Sieve element length varied with tree size, while sieve element radius, sieve pore radius, and the number of sieve areas per compound plate varied with sampling position. When data from all size classes were aggregated, all four variables followed a power‐law trend with distance from the top of the tree. The net effect of these ontogenetic scalings was to make total hydraulic sieve tube resistance independent of tree height from 0.5 to over 20 m.

    Conclusions

    Sieve element development responded to two pieces of information, tree size and distance from the apex, in a manner that conserved total sieve tube resistance across size classes. A further differentiated response between the phloem in the live crown and in the main stem is also suggested.

     
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