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  1. Abstract

    Southern California experienced unprecedented megadrought between 2012 and 2018. During this time,Malosma laurina, a chaparral species normally resilient to single‐year intense drought, developed extensive mortality exceeding 60% throughout low‐elevation coastal populations of the Santa Monica Mountains. We assessed the physiological mechanisms by which the advent of megadrought predisposedM. laurinato extensive shoot dieback and whole‐plant death. We found that hydraulic conductance of stem xylem (Ks, native) was reduced seven to 11‐fold in dieback adult and resprout branches, respectively. Staining of stem xylem vessels revealed that dieback plants experienced 68% solid‐blockage, explaining the reduction in water transport. Following Koch's postulates, persistent isolation of a microorganism in stem xylem of dieback plants but not healthy controls indicated that the causative agent of xylem blockage was an opportunistic endophytic fungus,Botryosphaeria dothidea. We inoculated healthyM. laurinasaplings with fungal isolates and compared hyphal elongation rates under well‐watered, water‐deficit, and carbon‐deficit treatments. Relative to controls, we found that both water deficit and carbon‐deficit increased hyphal extension rates and the incidence of shoot dieback.

     
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  2. Abstract Desiccation-tolerant (DT) organisms can lose nearly all their water without dying. Desiccation tolerance allows organisms to survive in a nearly completely dehydrated, dormant state. At the cellular level, sugars and proteins stabilize cellular components and protect them from oxidative damage. However, there are few studies of the dynamics and drivers of whole-plant recovery in vascular DT plants. In vascular DT plants, whole-plant desiccation recovery (resurrection) depends not only on cellular rehydration, but also on the recovery of organs with unequal access to water. In this study, in situ natural and artificial irrigation experiments revealed the dynamics of desiccation recovery in two DT fern species. Organ-specific irrigation experiments revealed that the entire plant resurrected when water was supplied to roots, but leaf hydration alone (foliar water uptake) was insufficient to rehydrate the stele and roots. In both species, pressure applied to petioles of excised desiccated fronds resurrected distal leaf tissue, while capillarity alone was insufficient to resurrect distal pinnules. Upon rehydration, sucrose levels in the rhizome and stele dropped dramatically as starch levels rose, consistent with the role of accumulated sucrose as a desiccation protectant. These findings provide insight into traits that facilitate desiccation recovery in dryland ferns associated with chaparral vegetation of southern California. 
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  3. Summary

    Desiccation‐tolerant (DT) plants can dry past −100 MPa and subsequently recover function upon rehydration. Vascular DT plants face the unique challenges of desiccating and rehydrating complex tissues without causing structural damage. However, these dynamics have not been studied in intact DT plants.

    We used high resolution micro‐computed tomography (microCT), light microscopy, and fluorescence microscopy to characterize the dynamics of tissue desiccation and rehydration in petioles (stipes) of intact DT ferns.

    During desiccation, xylem conduits in stipes embolized before cellular dehydration of living tissues within the vascular cylinder. During resurrection, the chlorenchyma and phloem within the stipe vascular cylinder rehydrated before xylem refilling. We identified unique stipe traits that may facilitate desiccation and resurrection of the vascular system, including xylem conduits containing pectin (which may confer flexibility and wettability); chloroplasts within the vascular cylinder; and an endodermal layer impregnated with hydrophobic substances that impede apoplastic leakage while facilitating the upward flow of water within the vascular cylinder.

    Resurrection ferns are a novel system for studying extreme dehydration recovery and embolism repair in the petioles of intact plants. The unique anatomical traits identified here may contribute to the spatial and temporal dynamics of water movement observed during desiccation and resurrection.

     
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  4. Summary

    Given increasing water deficits across numerous ecosystems world‐wide, it is urgent to understand the sequence of failure of leaf function during dehydration.

    We assessed dehydration‐induced losses of rehydration capacity and maximum quantum yield of the photosystemII(Fv/Fm) in the leaves of 10 diverse angiosperm species, and tested when these occurred relative to turgor loss, declines of stomatal conductancegs, and hydraulic conductanceKleaf, including xylem and outside xylem pathways for the same study plants. We resolved the sequences of relative water content and leaf water potential Ψleafthresholds of functional impairment.

    On average, losses of leaf rehydration capacity occurred at dehydration beyond 50% declines ofgs,Kleafand turgor loss point. Losses ofFv/Fmoccurred after much stronger dehydration and were not recovered with leaf rehydration. Across species, tissue dehydration thresholds were intercorrelated, suggesting trait co‐selection. Thresholds for each type of functional decline were much less variable across species in terms of relative water content than Ψleaf.

    The stomatal and leaf hydraulic systems show early functional declines before cell integrity is lost. Substantial damage to the photochemical apparatus occurs at extreme dehydration, after complete stomatal closure, and seems to be irreversible.

     
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