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Leaf shrinkage with dehydration has attracted attention for over 100 years, especially as it becomes visibly extreme during drought. However, little has been known of its correlation with physiology. Computer simulations of the leaf hydraulic system showed that a reduction of hydraulic conductance of the mesophyll pathways outside the xylem would cause a strong decline of leaf hydraulic conductance (Kleaf). For 14 diverse species, we tested the hypothesis that shrinkage during dehydration (i.e. in whole leaf, cell and airspace thickness, and leaf area) is associated with reduction in Kleaf at declining leaf water potential (Ψleaf). We tested hypotheses for the linkage of leaf shrinkage with structural and physiological water relations parameters, including modulus of elasticity, osmotic pressure at full turgor, turgor loss point (TLP), and cuticular conductance. Species originating from moist habitats showed substantial shrinkage during dehydration before reaching TLP, in contrast with species originating from dry habitats. Across species, the decline of Kleaf with mild dehydration (i.e. the initial slope of the Kleaf versus Ψleaf curve) correlated with the decline of leaf thickness (the slope of the leaf thickness versus Ψleaf curve), as expected based on predictions from computer simulations. Leaf thickness shrinkage before TLP correlated across species with lower modulus of elasticity and with less negative osmotic pressure at full turgor, as did leaf area shrinkage between full turgor and oven desiccation. These findings point to a role for leaf shrinkage in hydraulic decline during mild dehydration, with potential impacts on drought adaptation for cells and leaves, influencing plant ecological distributions.As leaves open their stomata to capture CO2 for photosynthesis, water is lost to transpiration, which needs to be replaced by flow through the hydraulic system. The leaf hydraulic system has two components, which act essentially in series: the pathways for water movement through the xylem from the petiole to leaf minor veins, and those through the living bundle sheath and mesophyll cells to the sites of evaporation (Tyree and Zimmermann, 2002; Sack et al., 2004; Sack and Holbrook, 2006). The decline in leaf hydraulic conductance (Kleaf) with dehydration may thus depend on both components. The importance of the xylem component is well established. Vein xylem embolism and cell collapse have been observed in dehydrating leaves (Salleo et al., 2001; Cochard et al., 2004a; Johnson et al., 2009), and computer modeling and experimental work showed that species with high major vein length per leaf area (VLA; i.e. for the first three vein-branching orders) were more resistant to hydraulic decline, providing more pathways around embolisms (Scoffoni et al., 2011). However, the physical impacts of dehydration on the extraxylem pathways have not been studied, even though in turgid leaves these pathways account for 26% to 88% of leaf hydraulic resistance (i.e. of 1/Kleaf), depending on species (Sack et al., 2003a; Cochard et al., 2004b). The aim of this study was to determine whether leaf shrinkage during dehydration relates to the decline of Kleaf as well as the structural determinants of leaf shrinkage.The shrinkage of leaves with dehydration has drawn attention for over 100 years. Leaves shrink in their area (Bogue, 1892; Gardner and Ehlig, 1965; Jones, 1973; Tang and Boyer, 2007; Blonder et al., 2012) and, considered in relative terms, even more strongly in their thickness (Fig. 1; Meidner, 1952; Gardner and Ehlig, 1965; Downey and Miller, 1971; Syvertsen and Levy, 1982; Saini and Rathore, 1983; Burquez, 1987; McBurney, 1992; Sancho-Knapik et al., 2010, 2011). Leaves fluctuate in thickness daily and seasonally according to transpiration (Kadoya et al., 1975; Tyree and Cameron, 1977; Fensom and Donald, 1982; Rozema et al., 1987; Ogaya and Peñuelas, 2006; Seelig et al., 2012). Indeed, the relation of leaf thickness to water status is so tight that using leaf thickness to guide irrigation has led to water savings of up to 45% (Seelig et al., 2012).Open in a separate windowFigure 1.Sketches of a fully turgid leaf (A) versus a strongly dehydrated leaf (B; drawings based on leaf cross sections of sunflower in Fellows and Boyer, 1978). Note the strong reduction in leaf thickness, cell thickness, and intercellular airspaces in the dehydrated leaf. Epidermal cells are shrunk in the dehydrated leaf, inducing whole-leaf area shrinkage. Note that this sketch represents shrinkage for a typical drought-sensitive species. Many species such as oaks (Quercus spp.) will experience less thickness shrinkage and an increase in intercellular airspace (see “Discussion”). [See online article for color version of this figure.]Previous studies of leaf shrinkage with progressive dehydration have tended to focus on single or few species. These studies showed that thickness declines with water status in two phases. Before the bulk leaf turgor loss point (TLP; leaf water potential [Ψleaf] at TLP) is reached, the slope of leaf thickness versus Ψleaf or relative water content (RWC) is shallower than past TLP for most species (Meidner, 1955, Kennedy and Booth, 1958, Burquez, 1987, McBurney, 1992, Sancho-Knapik et al., 2010, 2011). This is because before TLP, declining Ψleaf is strongly driven by declines in turgor pressure, which have a relatively low impact on cell and airspace volume, whereas past the TLP, declining Ψleaf depends only on solute concentration, which increases in inverse proportion as cell water volume declines while airspaces may shrink or expand (Tyree and Hammel, 1972, Sancho-Knapik et al., 2011). However, the steepness of the slope of leaf thickness versus Ψleaf before TLP seems to vary strongly across species (Meidner, 1955; Kennedy and Booth, 1958; Fellows and Boyer, 1978; Burquez, 1987; Colpitts and Coleman, 1997; Sancho-Knapik et al., 2010).A high leaf cell volume and turgor is crucial to physiological processes (Boyer, 1968; Lawlor and Cornic, 2002). Shrinkage may affect cell connectivity and water transport (Sancho-Knapik et al., 2011). However, no studies have tested for a possible relationship of leaf shrinkage with the decline of Kleaf during dehydration. Such an association would arise if, across species, shrinkage occurred simultaneously with vein xylem embolism or if tissue shrinkage led to declines in the extraxylem hydraulic conductance.To refine our hypotheses, we modified a computer model of the leaf hydraulic system (Cochard et al., 2004b; McKown et al., 2010; Scoffoni et al., 2011) to predict the impact of losses of xylem and extraxylem conductance on the response of Kleaf to dehydration. We characterized the degree of leaf shrinkage in thickness, in the thickness of cells and airspaces within the leaf, and in leaf area for 14 species diverse in phylogeny, leaf traits, and drought tolerance. We hypothesized that loss of extraxylem hydraulic conductance should have a greater impact on Kleaf at less negative water potentials when xylem tensions are too weak to trigger embolism and induce dramatic declines in Kleaf. We hypothesized that species with greater degrees of shrinkage before TLP would experience greater loss of Kleaf. Furthermore, we hypothesized that species from moist habitats would have greater degrees of shrinkage.For insight into the mechanisms and consequences of leaf shrinkage, we also investigated the relationships of 18 indices of leaf shrinkage with a wide range of aspects of leaf structure and composition, including gross morphology, leaf venation architecture, parameters of pressure-volume curves, and leaf water storage. We hypothesized that, across species, shrinkage in whole leaf, cell, and intercellular airspace thickness would be lower for species with greater allocation to structural rigidity and osmotic concentration, and thus shrinkage would be positively correlated with a lower modulus of elasticity (ε), less negative osmotic pressure at full turgor (πo), lower leaf mass per area (LMA), and lower leaf density. Additionally, we tested the longstanding hypothesis that species with higher major VLA and/or minor VLA (i.e. the fourth and higher vein-branching orders) would shrink less in area and/or thickness with dehydration (Gardner and Ehlig, 1965). Finally, we tested the ability of dehydrated leaves to recover in size with rehydration. We hypothesized that recovery would be greater for mildly than for strongly dehydrated leaves and that species with greater leaf shrinkage would be better able to recover from shrinkage.  相似文献   
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Plant trichomes play important protective functions and may have a major influence on leaf surface wettability. With the aim of gaining insight into trichome structure, composition, and function in relation to water-plant surface interactions, we analyzed the adaxial and abaxial leaf surface of holm oak (Quercus ilex) as a model. By measuring the leaf water potential 24 h after the deposition of water drops onto abaxial and adaxial surfaces, evidence for water penetration through the upper leaf side was gained in young and mature leaves. The structure and chemical composition of the abaxial (always present) and adaxial (occurring only in young leaves) trichomes were analyzed by various microscopic and analytical procedures. The adaxial surfaces were wettable and had a high degree of water drop adhesion in contrast to the highly unwettable and water-repellent abaxial holm oak leaf sides. The surface free energy and solubility parameter decreased with leaf age, with higher values determined for the adaxial sides. All holm oak leaf trichomes were covered with a cuticle. The abaxial trichomes were composed of 8% soluble waxes, 49% cutin, and 43% polysaccharides. For the adaxial side, it is concluded that trichomes and the scars after trichome shedding contribute to water uptake, while the abaxial leaf side is highly hydrophobic due to its high degree of pubescence and different trichome structure, composition, and density. Results are interpreted in terms of water-plant surface interactions, plant surface physical chemistry, and plant ecophysiology.Plant surfaces have an important protecting function against multiple biotic and abiotic stress factors (Riederer, 2006). They may, for example, limit the attack of insects (Eigenbrode and Jetter, 2002) or pathogenic fungi (Gniwotta et al., 2005; Łaźniewska et al., 2012), avoid damage caused by high intensities of UV and visible radiation (Reicosky and Hanover, 1978; Karabourniotis and Bormann, 1999), help to regulate leaf temperature (Ehleringer and Björkman, 1978; Ripley et al., 1999), and chiefly prevent plant organs from dehydration (Riederer and Schreiber, 2001).The epidermis of plants has been found to have a major degree of physical and chemical variability and may often contain specialized cells such as trichomes or stomata (Roth-Nebelsick et al., 2009; Javelle et al., 2011). Most aerial organs are covered with an extracellular and generally lipid-rich layer named the cuticle, which is typically composed of waxes embedded in (intracuticular waxes) or deposited on (epicuticular waxes) a biopolymer matrix of cutin, forming a network of cross-esterified hydroxy C16 and/or C18 fatty acids, and/or cutan, with variable amounts of polysaccharides and phenolics (Domínguez et al., 2011; Yeats and Rose, 2013). Different nano- and/or microscale levels of plant surface sculpturing have been observed by scanning electron microscopy (SEM), generally in relation to the topography of epicuticular waxes, cuticular folds, and epidermal cells (Koch and Barthlott, 2009). Such surface features together with their chemical composition (Khayet and Fernández, 2012) may lead to a high degree of roughness and hydrophobicity (Koch and Barthlott, 2009; Konrad et al., 2012). The interactions of plant surfaces with water have been addressed in some investigations (Brewer et al., 1991; Brewer and Smith, 1997; Pandey and Nagar, 2003; Hanba et al., 2004; Dietz et al., 2007; Holder, 2007a, 2007b; Fernández et al., 2011, 2014; Roth-Nebelsick et al., 2012; Wen et al., 2012; Urrego-Pereira et al., 2013) and are a topic of growing interest for plant ecophysiology (Helliker and Griffiths, 2007; Aryal and Neuner, 2010; Limm and Dawson, 2010; Kim and Lee, 2011; Berry and Smith, 2012; Berry et al., 2013; Rosado and Holder, 2013; Helliker, 2014). On the other hand, the mechanisms of foliar uptake of water and solutes by plant surfaces are still not fully understood (Fernández and Eichert, 2009; Burkhardt and Hunsche, 2013), but they may play an important ecophysiological role (Limm et al., 2009; Johnstone and Dawson, 2010; Adamec, 2013; Berry et al., 2014).The importance of trichomes and pubescent layers on water drop-plant surface interactions and on the subsequent potential water uptake into the organs has been analyzed in some investigations (Fahn, 1986; Brewer et al., 1991; Grammatikopoulos and Manetas, 1994; Brewer and Smith, 1997; Pierce et al., 2001; Kenzo et al., 2008; Fernández et al., 2011, 2014; Burrows et al., 2013). Trichomes are unicellular or multicellular and glandular or nonglandular appendages, which originate from epidermal cells only and develop outwards on the surface of plant organs (Werker, 2000). Nonglandular trichomes are categorized according to their morphology and exhibit a major variability in size, morphology, and function. On the other hand, glandular trichomes are classified by the secretory materials they excrete, accumulate, or absorb (Johnson, 1975; Werker, 2000; Wagner et al., 2004). Trichomes can be often found in xeromorphic leaves and in young organs (Fahn, 1986; Karabourniotis et al., 1995). The occurrence of protecting leaf trichomes has been also reported for Mediterranean species such as holm oak (Quercus ilex; Karabourniotis et al., 1995, 1998; Morales et al., 2002; Karioti et al., 2011; Camarero et al., 2012). There is limited information about the nature of the surface of trichomes, but they are also covered with a cuticle similarly to other epidermal cell types (Fernández et al., 2011, 2014).In this study and using holm oak as a model, we assessed, for the first time, the leaf surface-water relations of the abaxial (always pubescent) versus the adaxial (only pubescent in developing leaves and for a few months) surface, including their capacity to absorb surface-deposited water drops. Based on membrane science methodologies (Fernández et al., 2011; Khayet and Fernández, 2012) and following a new integrative approach, the chemical, physical, and anatomical properties of holm oak leaf surfaces and trichomes were analyzed, with the aim of addressing the following questions. Are young and mature adaxial and abaxial leaf surfaces capable of absorbing water deposited as drops on to the surfaces? Are young and mature abaxial and adaxial leaf surfaces similar in relation to their wettability, hydrophobicity, polarity, work of adhesion (Wa) for water, solubility parameter (δ), and surface free energy (γ)? What is the physical and chemical nature of the adaxial versus the abaxial trichomes, chiefly in relation to young leaves?  相似文献   
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Plasma lipidome is now increasingly recognized as a potentially important marker of chronic diseases, but the exact extent of its contribution to the interindividual phenotypic variability in family studies is unknown. Here, we used the rich data from the ongoing San Antonio Family Heart Study (SAFHS) and developed a novel statistical approach to quantify the independent and additive value of the plasma lipidome in explaining metabolic syndrome (MS) variability in Mexican American families recruited in the SAFHS. Our analytical approach included two preprocessing steps: principal components analysis of the high-resolution plasma lipidomics data and construction of a subject-subject lipidomic similarity matrix. We then used the Sequential Oligogenic Linkage Analysis Routines software to model the complex family relationships, lipidomic similarities, and other important covariates in a variance components framework. Our results suggested that even after accounting for the shared genetic influences, indicators of lipemic status (total serum cholesterol, TGs, and HDL cholesterol), and obesity, the plasma lipidome independently explained 22% of variability in the homeostatic model of assessment-insulin resistance trait and 16% to 22% variability in glucose, insulin, and waist circumference. Our results demonstrate that plasma lipidomic studies can additively contribute to an understanding of the interindividual variability in MS.  相似文献   
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In this study, we investigated the expression levels of host restriction factors in six untreated HIV-1-positive patients over the course of infection. We found that the host restriction factor gene expression profile consistently increased over time and was significantly associated with CD4+ T cell activation and viral load. Our data are among the first to demonstrate the dynamic nature of host restriction factors in vivo over time.  相似文献   
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Highly pathogenic avian influenza H5N1 virus clades 2.3.4, 2.3.2, and 7 are the dominant cocirculating H5N1 viruses in poultry in China. However, humans appear to be clinically susceptible mostly to the 2.3.4 virus clade. Here, we demonstrated that A549 cells and human macrophages infected with clade 2.3.4 viruses produced significantly more viruses than those infected with the other two clades. Likewise, clade 2.3.4-infected macrophages caused the most severe cellular damage and strongest proinflammatory response.  相似文献   
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