Leaf and stem physiological responses to summer and winter extremes of woody species across temperate ecosystems

© 2014 The Authors. Winter cold limits temperate plant performance, as does summer water stress in drought-prone ecosystems. The relative impact of seasonal extremes on plant performance has received considerable attention for individual systems. An integrated study compiling the existing literature was needed to identify overall trends. First, we conducted a meta-analysis of the impacts of summer and winter on ecophysiology for three woody plant functional types (winter deciduous angiosperms, evergreen angiosperms and conifers), including data for 210 records from 75 studies of ecosystems with and without summer drought across the temperate zone. Second, we tested predictions by conducting a case study in a drought-prone Mediterranean ecosystem subject to winter freezing. As indicators of physiological response of leaves and xylem to seasonal stress, we focused on stomatal conductance (gs), percent loss of stem xylem hydraulic conductivity (PLC) and photochemical efficiency of photosystem II (Fv/Fm). Our meta-analysis showed that in ecosystems without summer drought, gs was higher during summer than winter. By contrast, in drought-prone ecosystems many species maintained open stomata during winter, with potential strong consequences for plant carbon gain over the year. Further, PLC tended to increase and Fv/Fm to decrease from summer to winter for most functional types and ecosystems due to low temperatures. Overall, deciduous angiosperms were most sensitive to climatic stress. Leaf gas exchange and stem xylem hydraulics showed a coordinated seasonal response at ecosystems without summer drought. In our Mediterranean site subjected to winter freezing the species showed similar responses to those typically found for ecosystems without summer drought. We conclude that winter stress is most extreme for systems without summer drought and systems with summer drought and winter freezing, and less extreme for drought-prone systems without freezing. In all cases the evergreen species show less pronounced seasonal responses in both leaves and stems than deciduous species.Th is research was supported by the Spanish Ministry of Economy and Competitiveness with the grants FPI (CGL2007-66066-C04-02), Consolider Montes (CSD2008 00040) and VULGLO (CGL2010 22180 C03 03), and by the Community of Madrid grant REMEDINAL 2 (CM S2009 AMB 1783) and National Science Foundation Grant no. 0546784.Peer Reviewe

Resolving Australian analogs for an Eocene Patagonian paleorainforest using leaf size and floristics

• Premise of the study: The diverse early Eocene flora from Laguna del Hunco (LH) in Patagonia, Argentina has many nearest living relatives (NLRs) in Australasia but few in South America, indicating the differential survival of an ancient, trans‐Antarctic rainforest biome. To better understand this significant biogeographic pattern, we used detailed comparisons of leaf size and floristics to quantify the legacy of LH across a large network of Australian rainforest‐plot assemblages. • Methods: We applied vein scaling, a new method for estimating the original areas of fragmented leaves. We then compared leaf size and floristics at LH with living Australian assemblages and tabulated the climates of those where NLRs occur, along with additional data on climatic ranges of “ex‐Australian” NLRs that survive outside of Australia. • Key results: Vein scaling estimated areas as accurately as leaf‐size classes. Applying vein scaling to fossil fragments increased the grand mean area of LH by 450 mm2, recovering more originally large leaves. The paleoflora has a majority of microphyll leaves, comparable to leaf litter in subtropical Australian forests, which also have the greatest floristic similarity to LH. Tropical Australian assemblages also share many taxa with LH, and ex‐Australian NLRs mostly inhabit cool, wet montane habitats no longer present in Australia. • Conclusions: Vein scaling is valuable for improving the resolution of fossil leaf‐size distributions by including fragmented specimens. The legacy of LH is evident not only in subtropical and tropical Australia but also in tropical montane Australasia and Southeast Asia, reflecting the disparate histories of surviving Gondwanan lineages.Fil: Merkhofer, Lisa. State University of Pennsylvania; Estados UnidosFil: Wilf, Peter. State University of Pennsylvania; Estados UnidosFil: Haas, M. Tyler. State University of Pennsylvania; Estados UnidosFil: Kooyman, Robert M.. Macquarie University; AustraliaFil: Sack, Lawren. University of California at Los Angeles; Estados UnidosFil: Scoffoni, Christine. University of California at Los Angeles; Estados UnidosFil: Cúneo, Néstor Rubén. Museo Paleontológico Egidio Feruglio; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentin

Leaf vein length per unit area is not intrinsically dependent on image magnification: avoiding measurement artifacts for accuracy and precision.

Leaf vein length per unit leaf area (VLA; also known as vein density) is an important determinant of water and sugar transport, photosynthetic function, and biomechanical support. A range of software methods are in use to visualize and measure vein systems in cleared leaf images; typically, users locate veins by digital tracing, but recent articles introduced software by which users can locate veins using thresholding (i.e. based on the contrasting of veins in the image). Based on the use of this method, a recent study argued against the existence of a fixed VLA value for a given leaf, proposing instead that VLA increases with the magnification of the image due to intrinsic properties of the vein system, and recommended that future measurements use a common, low image magnification for measurements. We tested these claims with new measurements using the software LEAFGUI in comparison with digital tracing using ImageJ software. We found that the apparent increase of VLA with magnification was an artifact of (1) using low-quality and low-magnification images and (2) errors in the algorithms of LEAFGUI. Given the use of images of sufficient magnification and quality, and analysis with error-free software, the VLA can be measured precisely and accurately. These findings point to important principles for improving the quantity and quality of important information gathered from leaf vein systems

Developmental and biophysical determinants of grass leaf size worldwide

One of the most notable ecological trends—described more than 2,300 years ago by Theophrastus—is the association of small leaves with dry and cold climates, which has recently been recognized for eudicotyledonous plants at a global scale. For eudicotyledons, this pattern has been attributed to the fact that small leaves have a thinner boundary layer that helps to avoid extreme leaf temperatures and their leaf development results in vein traits that improve water transport under cold or dry climates. However, the global distribution of leaf size and its adaptive basis have not been tested in the grasses, which represent a diverse lineage that is distinct in leaf morphology and that contributes 33% of terrestrial primary productivity (including the bulk of crop production). Here we demonstrate that grasses have shorter and narrower leaves under colder and drier climates worldwide. We show that small grass leaves have thermal advantages and vein development that contrast with those of eudicotyledons, but that also explain the abundance of small leaves in cold and dry climates. The worldwide distribution of leaf size in grasses exemplifies how biophysical and developmental processes result in convergence across major lineages in adaptation to climate globally, and highlights the importance of leaf size and venation architecture for grass performance in past, present and future ecosystems

Deciduous and evergreen oaks show contrasting adaptive responses in leaf mass per area across environments

• Increases in leaf mass per area (LMA) are commonly observed in response to environmental stresses and are achieved through increases in leaf thickness and/or leaf density. Here, we investigated how the two underlying components of LMA differ in relation to species native climates and phylogeny, across deciduous and evergreen species. • Using a phylogenetic approach, we quantified anatomical, compositional and climatic variables from 40 deciduous and 45 evergreen Quercus species from across the Northern Hemisphere growing in a common garden. • Deciduous from shorter growing seasons tended to have leaves with lower LMA and leaf thickness than those from longer growing seasons, while the opposite pattern was found for evergreens. For both habits, LMA and thickness increased in arid environments. However, this shift was associated with increased leaf density in evergreens but reduced density in deciduous species. • Deciduous and evergreen oaks showed fundamental leaf morphological differences that revealed a diverse adaptive response. While LMA in deciduous may diversified in tight coordination with thickness mainly modulated by aridity, diversification of LMA within evergreens appears dependent on the infrageneric group, with diversification in leaf thickness modulated by both aridity and cold, while diversification in leaf density only modulated by aridity.Publishe

The handbook for standardized field and laboratory measurements in terrestrial climate change experiments and observational studies (ClimEx)

1. Climate change is a world‐wide threat to biodiversity and ecosystem structure, functioning and services. To understand the underlying drivers and mechanisms, and to predict the consequences for nature and people, we urgently need better understanding of the direction and magnitude of climate change impacts across the soil–plant–atmosphere continuum. An increasing number of climate change studies are creating new opportunities for meaningful and high‐quality generalizations and improved process understanding. However, significant challenges exist related to data availability and/or compatibility across studies, compromising opportunities for data re‐use, synthesis and upscaling. Many of these challenges relate to a lack of an established ‘best practice’ for measuring key impacts and responses. This restrains our current understanding of complex processes and mechanisms in terrestrial ecosystems related to climate change. 2. To overcome these challenges, we collected best‐practice methods emerging from major ecological research networks and experiments, as synthesized by 115 experts from across a wide range of scientific disciplines. Our handbook contains guidance on the selection of response variables for different purposes, protocols for standardized measurements of 66 such response variables and advice on data management. Specifically, we recommend a minimum subset of variables that should be collected in all climate change studies to allow data re‐use and synthesis, and give guidance on additional variables critical for different types of synthesis and upscaling. The goal of this community effort is to facilitate awareness of the importance and broader application of standardized methods to promote data re‐use, availability, compatibility and transparency. We envision improved research practices that will increase returns on investments in individual research projects, facilitate second‐order research outputs and create opportunities for collaboration across scientific communities. Ultimately, this should significantly improve the quality and impact of the science, which is required to fulfil society's needs in a changing world

Leaf hydraulics and evolution

There has been increasing worldwide recognition of the importance of hydraulic physiology--the transport of water through the plant--in explaining plant growth and drought tolerance. By combining physiology and anatomy within an evolutionary framework, we can discover the mechanisms underlying species differences in hydraulic function, especially those of the leaf, the central organ in plant metabolism. I refined and developed new methods to investigate leaf water transport and its decline during drought, focusing on a critical measure of the capacity for water movement (leaf hydraulic conductance, Kleaf). I found that species most tolerant of Kleaf decline had small leaves with dense major veins, providing pathways for the water to bypass embolized conduits during drought giving a new, direct explanation to the fact that species of dry areas have small leaves. I also developed a new method to investigate the role of leaf shrinkage on water movement. As leaves shrink with dehydration, mesophyll cells lose connectivity, physically impacting water movement outside the xylem. I found that species most sensitive to Kleaf decline were those with strongest shrinkage in thickness. I then developed a new method to measure xylem hydraulic decline in leaves to test for a possible artifact of cutting leaf petioles under tension while under water. Such artifact has been recently found to occur in stems, and has put into question measurements of Kleaf. Across four diverse species, I found no sign of such an artifact in leaves, likely due to the lesser mechanical stress imposed when cutting a petiole vs. stem. Finally, I took an evolutionary perspective. I quantified the anatomical and physiological plasticity in leaves of six species of endemic Hawaiian lobeliads grown under different light regimes and found a high degree of plasticity in Kleaf with light, relating to leaf anatomical changes. Across 30 species of Viburnum I have identified the evolutionary shifts of leaf anatomy, water transport and drought tolerance. This work provides new techniques, clarity and applications toward understanding leaf water transport and its role in plant performance and drought tolerance, with applications for ecology, paleobiology and the conservation of species and ecosystems

Why are leaves hydraulically vulnerable?

As plant tissues dehydrate, water transport efficiency declines, a process typically attributed to air obstruction (embolism) in the xylem. Trifiló et al. (pages 5029-5039) dissect leaf hydraulic vulnerability and show that both xylem and living tissues may be important. If confirmed and clarified, an important role for outsidexylem hydraulic decline will change our understanding of how plants transport water and control biosphere carbon and water fluxes