Showing papers in "New Phytologist in 2013"

Biosynthesis, function and metabolic engineering of plant volatile organic compounds.

Abstract: Plants synthesize an amazing diversity of volatile organic compounds (VOCs) that facilitate interactions with their environment, from attracting pollinators and seed dispersers to protecting themselves from pathogens, parasites and herbivores. Recent progress in -omics technologies resulted in the isolation of genes encoding enzymes responsible for the biosynthesis of many volatiles and contributed to our understanding of regulatory mechanisms involved in VOC formation. In this review, we largely focus on the biosynthesis and regulation of plant volatiles, the involvement of floral volatiles in plant reproduction as well as their contribution to plant biodiversity and applications in agriculture via crop-pollinator interactions. In addition, metabolic engineering approaches for both the improvement of plant defense and pollinator attraction are discussed in light of methodological constraints and ecological complications that limit the transition of crops with modified volatile profiles from research laboratories to real-world implementation.

Competition between roots and microorganisms for nitrogen: mechanisms and ecological relevance

Abstract: Demand of all living organisms on the same nutrients forms the basis for interspecific competition between plants and microorganisms in soils. This competition is especially strong in the rhizosphere. To evaluate competitive and mutualistic interactions between plants and microorganisms and to analyse ecological consequences of these interactions, we analysed 424 data pairs from 41 (15)N-labelling studies that investigated (15)N redistribution between roots and microorganisms. Calculated Michaelis-Menten kinetics based on K(m) (Michaelis constant) and V(max) (maximum uptake capacity) values from 77 studies on the uptake of nitrate, ammonia, and amino acids by roots and microorganisms clearly showed that, shortly after nitrogen (N) mobilization from soil organic matter and litter, microorganisms take up most N. Lower K(m) values of microorganisms suggest that they are especially efficient at low N concentrations, but can also acquire more N at higher N concentrations (V(max)) compared with roots. Because of the unidirectional flow of nutrients from soil to roots, plants are the winners for N acquisition in the long run. Therefore, despite strong competition between roots and microorganisms for N, a temporal niche differentiation reflecting their generation times leads to mutualistic relationships in the rhizosphere. This temporal niche differentiation is highly relevant ecologically because it: protects ecosystems from N losses by leaching during periods of slow or no root uptake; continuously provides roots with available N according to plant demand; and contributes to the evolutionary development of mutualistic interactions between roots and microorganisms.

Fungal community analysis by high-throughput sequencing of amplified markers - a user's guide

Abstract: Summary Novel high-throughput sequencing methods outperform earlier approaches in terms of resolution and magnitude. They enable identification and relative quantification of community members and offer new insights into fungal community ecology. These methods are currently taking over as the primary tool to assess fungal communities of plant-associated endophytes, pathogens, and mycorrhizal symbionts, as well as free-living saprotrophs. Taking advantage of the collective experience of six research groups, we here review the different stages involved in fungal community analysis, from field sampling via laboratory procedures to bioinformatics and data interpretation. We discuss potential pitfalls, alternatives, and solutions. Highlighted topics are challenges involved in: obtaining representative DNA/RNA samples and replicates that encompass the targeted variation in community composition, selection of marker regions and primers, options for amplification and multiplexing, handling of sequencing errors, and taxonomic identification. Without awareness of methodological biases, limitations of markers, and bioinformatics challenges, large-scale sequencing projects risk yielding artificial results and misleading conclusions.

APETALA2/Ethylene Responsive Factor (AP2/ERF) transcription factors: mediators of stress responses and developmental programs

Abstract: Transcription factors belonging to the APETALA2/Ethylene Responsive Factor (AP2/ERF) family are conservatively widespread in the plant kingdom. These regulatory proteins are involved in the control of primary and secondary metabolism, growth and developmental programs, as well as responses to environmental stimuli. Due to their plasticity and to the specificity of individual members of this family, AP2/ERF transcription factors represent valuable targets for genetic engineering and breeding of crops. In this review, we integrate the evidence collected from functional and structural studies to describe their different mechanisms of action and the regulatory pathways that affect their activity.

The mycorrhizal-associated nutrient economy: a new framework for predicting carbon-nutrient couplings in temperate forests.

Abstract: Summary Understanding the context dependence of ecosystem responses to global changes requires the development of new conceptual frameworks. Here we propose a framework for considering how tree species and their mycorrhizal associates differentially couple carbon (C) and nutrient cyclesintemperateforests.Giventhattreespeciespredominantlyassociatewithasingletypeof mycorrhizalfungi(arbuscularmycorrhizal(AM)fungiorectomycorrhizal(ECM)fungi),andthat the two types of fungi differ in their modes of nutrient acquisition, we hypothesize that the abundance of AM and ECM trees in a plot, stand, or region may provide an integrated index of biogeochemical transformations relevant to C cycling and nutrient retention. First, we describe how forest plots dominated by AM tree species have nutrient economies that differ in their C– nutrientcouplingsfromthoseinplotsdominatedbyECMtrees.Secondly,wedemonstratehow the relative abundance of AM and ECM trees can be used to estimate nutrient dynamics across the landscape. Finally, we describe how our framework can be used to generate testable hypotheses about forest responses to global change factors, and how these dynamics can be used to develop better representations of plant–soil feedbacks and nutrient constraints on productivity in ecosystem and earth system models.

Resprouting as a key functional trait: how buds, protection and resources drive persistence after fire

Abstract: Resprouting as a response to disturbance is now widely recognized as a key functional trait among woody plants and as the basis for the persistence niche. However, the underlying mechanisms that define resprouting responses to disturbance are poorly conceptualized. Resprouting ability is constrained by the interaction of the disturbance regime that depletes the buds and resources needed to fund resprouting, and the environment that drives growth and resource allocation. We develop a buds-protection-resources (BPR) framework for understanding resprouting in fire-prone ecosystems, based on bud bank location, bud protection, and how buds are resourced. Using this framework we go beyond earlier emphases on basal resprouting and highlight the importance of apical, epicormic and below-ground resprouting to the persistence niche. The BPR framework provides insights into: resprouting typologies that include both fire resisters (i.e. survive fire but do not resprout) and fire resprouters; the methods by which buds escape fire effects, such as thick bark; and the predictability of community assembly of resprouting types in relation to site productivity, disturbance regime and competition. Furthermore, predicting the consequences of global change is enhanced by the BPR framework because it potentially forecasts the retention or loss of above-ground biomass.

Leaf venation: structure, function, development, evolution, ecology and applications in the past, present and future

Abstract: The design and function of leaf venation are important to plant performance, with key implications for the distribution and productivity of ecosystems, and applications in paleobiology, agriculture and technology. We synthesize classical concepts and the recent literature on a wide range of aspects of leaf venation. We describe 10 major structural features that contribute to multiple key functions, and scale up to leaf and plant performance. We describe the development and plasticity of leaf venation and its adaptation across environments globally, and a new global data compilation indicating trends relating vein length per unit area to climate, growth form and habitat worldwide. We synthesize the evolution of vein traits in the major plant lineages throughout paleohistory, highlighting the multiple origins of individual traits. We summarize the strikingly diverse current applications of leaf vein research in multiple fields of science and industry. A unified core understanding will enable an increasing range of plant biologists to incorporate leaf venation into their research.

Environmental and physiological determinants of carbon isotope discrimination in terrestrial plants

Abstract: Summary Stable carbon isotope ratios (d 13 C) of terrestrial plants are employed across a diverse range of applicationsinenvironmentalandplantsciences;however,thekindofinformationthatisdesired fromthed 13 Csignaloftendiffers.Attheextremes,itrangesbetweenpurely environmentaland purely biological. Here, we review environmental drivers of variation in carbon isotope discrimination (D) in terrestrial plants, and the biological processes that can either damp or amplifytheresponse.ForC3plants,whereDisprimarilycontrolledbytheratioofintercellularto ambient CO2 concentrations (ci/ca), coordination between stomatal conductance and photo- synthesisandleafareaadjustmenttendstoconstrainthepotentialenvironmentallydrivenrange of D. For C4 plants, variation in bundle-sheath leakiness to CO2 can either damp or amplify the effects of ci/ca on D. For plants with crassulacean acid metabolism (CAM), D varies over a relatively largerange asafunctionoftheproportion ofdaytimetonight-time CO2fixation. This range can be substantially broadened by environmental effects onD when carbon uptake takes place primarily during the day. The effective use of D across its full range of applications will require a holistic view of the interplay between environmental control and physiological modulation of the environmental signal.

Biogeographical patterns and determinants of invasion by forest pathogens in Europe

Abstract: A large database of invasive forest pathogens (IFPs) was developed to investigate the patterns and determinants of invasion in Europe. Detailed taxonomic and biological information on the invasive species was combined with country-specific data on land use, climate, and the time since invasion to identify the determinants of invasiveness, and to differentiate the class of environments which share territorial and climate features associated with a susceptibility to invasion. IFPs increased exponentially in the last four decades. Until 1919, IFPs already present moved across Europe. Then, new IFPs were introduced mainly from North America, and recently from Asia. Hybrid pathogens also appeared. Countries with a wider range of environments, higher human impact or international trade hosted more IFPs. Rainfall influenced the diffusion rates. Environmental conditions of the new and original ranges and systematic and ecological attributes affected invasiveness. Further spread of established IFPs is expected in countries that have experienced commercial isolation in the recent past. Densely populated countries with high environmental diversity may be the weakest links in attempts to prevent new arrivals. Tight coordination of actions against new arrivals is needed. Eradication seems impossible, and prevention seems the only reliable measure, although this will be difficult in the face of global mobility.

Short‐chain chitin oligomers from arbuscular mycorrhizal fungi trigger nuclear Ca2+ spiking in Medicago truncatula roots and their production is enhanced by strigolactone

Abstract: The primary objective of this study was to identify the molecular signals present in arbuscular mycorrhizal (AM) germinated spore exudates (GSEs) responsible for activating nuclear Ca(2+) spiking in the Medicago truncatula root epidermis. Medicago truncatula root organ cultures (ROCs) expressing a nuclear-localized cameleon reporter were used as a bioassay to detect AM-associated Ca(2+) spiking responses and LC-MS to characterize targeted molecules in GSEs. This approach has revealed that short-chain chitin oligomers (COs) can mimic AM GSE-elicited Ca(2+) spiking, with maximum activity observed for CO4 and CO5. This spiking response is dependent on genes of the common SYM signalling pathway (DMI1/DMI2) but not on NFP, the putative Sinorhizobium meliloti Nod factor receptor. A major increase in the CO4/5 concentration in fungal exudates is observed when Rhizophagus irregularis spores are germinated in the presence of the synthetic strigolactone analogue GR24. By comparison with COs, both sulphated and nonsulphated Myc lipochito-oligosaccharides (LCOs) are less efficient elicitors of Ca(2+) spiking in M. truncatula ROCs. We propose that short-chain COs secreted by AM fungi are part of a molecular exchange with the host plant and that their perception in the epidermis leads to the activation of a SYM-dependent signalling pathway involved in the initial stages of fungal root colonization.

Next‐generation dynamic global vegetation models: learning from community ecology

Abstract: ummary Dynamic global vegetation models (DGVMs) are powerful tools to project past, current and future vegetation patterns and associated biogeochemical cycles. However, most models are limited by how they define vegetation and by their simplistic representation of competition. We discuss how concepts from community assembly theory and coexistence theory can help to improve vegetation models. We further present a trait- and individual-based vegetation model (aDGVM2) that allows individual plants to adopt a unique combination of trait values. These traits define how individual plants grow and compete. A genetic optimization algorithm is used to simulate trait inheritance and reproductive isolation between individuals. These model properties allow the assembly of plant communities that are adapted to a site's biotic and abiotic conditions. The aDGVM2 simulates how environmental conditions influence the trait spectra of plant communities; that fire selects for traits that enhance fire protection and reduces trait diversity; and the emergence of life-history strategies that are suggestive of colonization–competition trade-offs. The aDGVM2 deals with functional diversity and competition fundamentally differently from current DGVMs. This approach may yield novel insights as to how vegetation may respond to climate change and we believe it could foster collaborations between functional plant biologists and vegetation modellers.

Senescence, ageing and death of the whole plant

Abstract: 'Summary' 696 I. 'Endgame theory' 697 II. 'Terms and conditions' 697 III. 'Juvenility and maturity' 699 IV. 'Sources and sinks' 700 V. 'Monocarpy and polycarpy' 703 VI. 'Longevity and ageing' 704 VII. 'Last words' 707 'Acknowledgements' 708 References 708 Summary This review considers the relationship between the lifespan of an individual plant and the longevity of its component cells, tissues and organs. It begins by defining the terms senescence, growth, development, turnover, ageing, death and program. Genetic and epigenetic mechanisms regulating phase change from juvenility to maturity influence directly the capacity for responding to senescence signals and factors determining reproduction-related patterns of deteriorative ageing and death. Senescence is responsive to communication between sources and sinks in which sugar signalling and hormonal regulation play central roles. Monocarpy and polycarpy represent contrasting outcomes of the balance between the determinacy of apical meristems and source–sink cross-talk. Even extremely long-lived perennials sustain a high degree of meristem integrity. Factors associated with deteriorative ageing in animals, such as somatic mutation, telomere attrition and the costs of repair and maintenance, do not seem to be particularly significant for plant lifespan, but autophagy-related regulatory networks integrated with nutrient signalling may have a part to play. Size is an important influence on physiological function and fitness of old trees. Self-control of modular structure allows trees to sustain viability over prolonged lifespans. Different turnover patterns of structural modules can account for the range of plant life histories and longevities.

A meta-analysis of experimental warming effects on terrestrial nitrogen pools and dynamics.

Abstract: Summary � Global warming may have profound effects on terrestrial ecosystems. However, a comprehensive evaluation of the effects of warming on ecosystem nitrogen (N) pools and dynamics is not available. � Here, we compiled data of 528 observations from 51 papers and carried out a meta-analysis of experimental warming effects on 13 variables related to terrestrial N pools and dynamics. � We found that, on average, net N mineralization and net nitrification rate were increased by 52.2 and 32.2%, respectively, under experimental warming treatment. N pools were also increased by warming, although the magnitude of this increase was less than that of N fluxes. Soil microbial N and N immobilization were not changed by warming, probably because microbes are limited by carbon sources. Grassland and shrubland/heathland were less responsive to warming than forest, probably because the reduction of soil moisture by warming offset the temperature effect in these areas. Soil heating cable and all-day treatment appeared to be the most effective method on N cycling among all treatment methods. � Results of this meta-analysis are useful for better understanding the response of N cycling to global warming and the underlying mechanism of warming effects on plants and ecosystem functions.

Drought response strategies define the relative contributions of hydraulic dysfunction and carbohydrate depletion during tree mortality.

Abstract: Summary Plant survival during drought requires adequate hydration in living tissues and carbohydrate reserves for maintenance and recovery. We hypothesized that tree growth and hydraulic strategy determines the intensity and duration of the ‘physiological drought’, thereby affecting the relative contributions of loss of hydraulic function and carbohydrate depletion during mortality. We compared patterns in growth rate, water relations, gas exchange and carbohydrate dynamics in three tree species subjected to prolonged drought. Two Eucalyptus species (E. globulus, E. smithii) exhibited high growth rates and water-use resulting in rapid declines in water status and hydraulic conductance. In contrast, conservative growth and water relations in Pinus radiata resulted in longer periods of negative carbon balance and significant depletion of stored carbohydrates in all organs. The ongoing demand for carbohydrates from sustained respiration highlighted the role that duration of drought plays in facilitating carbohydrate consumption. Two drought strategies were revealed, differentiated by plant regulation of water status: plants maximized gas exchange, but were exposed to low water potentials and rapid hydraulic dysfunction; and tight regulation of gas exchange at the cost of carbohydrate depletion. These findings provide evidence for a relationship between hydraulic regulation of water status and carbohydrate depletion during terminal drought.

Identification and characterization of MYB‐bHLH‐WD40 regulatory complexes controlling proanthocyanidin biosynthesis in strawberry (Fragaria × ananassa) fruits

Abstract: Summary Strawberry (Fragaria × ananassa) fruits contain high concentrations of flavonoids. In unripe strawberries, the flavonoids are mainly represented by proanthocyanidins (PAs), while in ripe fruits the red-coloured anthocyanins also accumulate. Most of the structural genes leading to PA biosynthesis in strawberry have been characterized, but no information is available on their transcriptional regulation. In Arabidopsis thaliana the expression of the PA biosynthetic genes is specifically induced by a ternary protein complex, composed of AtTT2 (AtMYB123), AtTT8 (AtbHLH042) and AtTTG1 (WD40-repeat protein). A strategy combining yeast-two-hybrid screening and agglomerative hierarchical clustering of transcriptomic and metabolomic data was undertaken to identify strawberry PA regulators. Among the candidate genes isolated, four were similar to AtTT2, AtTT8 and AtTTG1 (FaMYB9/FaMYB11, FabHLH3 and FaTTG1, respectively) and two encode putative negative regulators (FaMYB5 and FabHLH3∆). Interestingly, FaMYB9/FaMYB11, FabHLH3 and FaTTG1 were found to complement the tt2-1, tt8-3 and ttg1-1 transparent testa mutants, respectively. In addition, they interacted in yeast and activated the Arabidopsis BANYULS (anthocyanidin reductase) gene promoter when coexpressed in Physcomitrella patens protoplasts. Taken together, these results demonstrated that FaMYB9/FaMYB11, FabHLH3 and FaTTG1 are the respective functional homologues of AtTT2, AtTT8 and AtTTG1, providing new tools for modifying PA content and strawberry fruit quality.

Evaluating theories of drought‐induced vegetation mortality using a multimodel–experiment framework

Abstract: 'Summary' 305 I. 'Background' 305 II. 'Model–experiment approach' 306 III. 'Simulations of hydraulic failure and carbon starvation' 310 IV. 'On thresholds vs duration of stress as drivers of mortality' 311 V. 'Interdependence of hydraulic failure and carbon starvation' 314 VI. 'Next-generation, traditional, and empirical models' 316 VII. 'A path forward' 317 VIII. 'Conclusions' 318 'Acknowledgements' 318 References 318 Summary Model–data comparisons of plant physiological processes provide an understanding of mechanisms underlying vegetation responses to climate. We simulated the physiology of a pinon pine–juniper woodland (Pinus edulis–Juniperus monosperma) that experienced mortality during a 5 yr precipitation-reduction experiment, allowing a framework with which to examine our knowledge of drought-induced tree mortality. We used six models designed for scales ranging from individual plants to a global level, all containing state-of-the-art representations of the internal hydraulic and carbohydrate dynamics of woody plants. Despite the large range of model structures, tuning, and parameterization employed, all simulations predicted hydraulic failure and carbon starvation processes co-occurring in dying trees of both species, with the time spent with severe hydraulic failure and carbon starvation, rather than absolute thresholds per se, being a better predictor of impending mortality. Model and empirical data suggest that limited carbon and water exchanges at stomatal, phloem, and below-ground interfaces were associated with mortality of both species. The model–data comparison suggests that the introduction of a mechanistic process into physiology-based models provides equal or improved predictive power over traditional process-model or empirical thresholds. Both biophysical and empirical modeling approaches are useful in understanding processes, particularly when the models fail, because they reveal mechanisms that are likely to underlie mortality. We suggest that for some ecosystems, integration of mechanistic pathogen models into current vegetation models, and evaluation against observations, could result in a breakthrough capability to simulate vegetation dynamics.

Sensitivity of plants to changing atmospheric CO2 concentration: From the geological past to the next century

Abstract: 'Summary' 1077 I. 'Introduction' 1078 II. 'Atmospheric CO2 concentrations through time' 1079 III. 'Plant sensitivity to CO2 at geological timescales' 1080 IV. 'Plant sensitivity to CO2 over the last 200 yr' 1081 V. 'Plant sensitivity to long-term experimental manipulation of CO2' 1084 VI. 'Simple formulation of stomatal conductance in land surface models for simulating long-term CO2 response' 1087 VII. 'Conclusions' 1088 'Acknowledgements' 1089 References 1089 Summary The rate of CO2 assimilation by plants is directly influenced by the concentration of CO2 in the atmosphere, ca. As an environmental variable, ca also has a unique global and historic significance. Although relatively stable and uniform in the short term, global ca has varied substantially on the timescale of thousands to millions of years, and currently is increasing at seemingly an unprecedented rate. This may exert profound impacts on both climate and plant function. Here we utilise extensive datasets and models to develop an integrated, multi-scale assessment of the impact of changing ca on plant carbon dioxide uptake and water use. We find that, overall, the sensitivity of plants to rising or falling ca is qualitatively similar across all scales considered. It is characterised by an adaptive feedback response that tends to maintain 1 − ci/ca, the relative gradient for CO2 diffusion into the leaf, relatively constant. This is achieved through predictable adjustments to stomatal anatomy and chloroplast biochemistry. Importantly, the long-term response to changing ca can be described by simple equations rooted in the formulation of more commonly studied short-term responses.

Cospeciation vs host-shift speciation: methods for testing, evidence from natural associations and relation to coevolution

Abstract: 'Summary' 347 I. 'Introduction' 348 II. 'Origin of the cospeciation concept' 349 III. 'Theoretical framework and methods for testing for cospeciation' 349 IV. 'Studies of natural associations reveal the prevalence of host shifts' 355 V. 'Relationship between host–symbiont coevolution and symbiont speciation' 378 VI. 'Conclusion' 381 'Acknowledgements' 381 References 381 Glossary 379 Summary Hosts and their symbionts are involved in intimate physiological and ecological interactions. The impact of these interactions on the evolution of each partner depends on the time-scale considered. Short-term dynamics – ‘coevolution’ in the narrow sense – has been reviewed elsewhere. We focus here on the long-term evolutionary dynamics of cospeciation and speciation following host shifts. Whether hosts and their symbionts speciate in parallel, by cospeciation, or through host shifts, is a key issue in host–symbiont evolution. In this review, we first outline approaches to compare divergence between pairwise associated groups of species, their advantages and pitfalls. We then consider recent insights into the long-term evolution of host–parasite and host–mutualist associations by critically reviewing the literature. We show that convincing cases of cospeciation are rare (7%) and that cophylogenetic methods overestimate the occurrence of such events. Finally, we examine the relationships between short-term coevolutionary dynamics and long-term patterns of diversification in host–symbiont associations. We review theoretical and experimental studies showing that short-term dynamics can foster parasite specialization, but that these events can occur following host shifts and do not necessarily involve cospeciation. Overall, there is now substantial evidence to suggest that coevolutionary dynamics of hosts and parasites do not favor long-term cospeciation.

Seasonal dynamics and age of stemwood nonstructural carbohydrates in temperate forest trees

Abstract: Nonstructural carbohydrate reserves support tree metabolism and growth when current photosynthates are insufficient, offering resilience in times of stress We monitored stemwood nonstructural carbohydrate (starch and sugars) concentrations of the dominant tree species at three sites in the northeastern United States We estimated the mean age of the starch and sugars in a subset of trees using the radiocarbon ((14) C) bomb spike With these data, we then tested different carbon (C) allocation schemes in a process-based model of forest C cycling We found that the nonstructural carbohydrates are both highly dynamic and about a decade old Seasonal dynamics in starch (two to four times higher in the growing season, lower in the dormant season) mirrored those of sugars Radiocarbon-based estimates indicated that the mean age of the starch and sugars in red maple (Acer rubrum) was 7-14 yr A two-pool (fast and slow cycling reserves) model structure gave reasonable estimates of the size and mean residence time of the total NSC pool, and greatly improved model predictions of interannual variability in woody biomass increment, compared with zero- or one-pool structures used in the majority of existing models This highlights the importance of nonstructural carbohydrates in the context of forest ecosystem carbon cycling

Mycorrhizal fungal establishment in agricultural soils: factors determining inoculation success.

Abstract: Soil biota provide a number of key ecological services to natural and agricultural ecosystems. Increasingly, inoculation of soils with beneficial soil biota is being considered as a tool to enhance plant productivity and sustainability of agricultural ecosystems. However, one important bottleneck is the establishment of viable microbial populations that can persist over multiple seasons. Here, we explore the factors responsible for establishment of the beneficial soil fungi, arbuscular mycorrhizal fungi (AMF), which can enhance the yield of a wide range of agricultural crops. We evaluate field application potential and discuss ecological and evolutionary factors responsible for application success. We identify three factors that determine inoculation success and AM fungal persistence in soils: species compatibility (can the introduced species thrive under the imposed circumstances?); field carrying capacity (the habitat niche available to AMF); and priority effects (the influence of timing and competition on the establishment of alternative stable communities). We explore how these factors can be employed for establishment and persistence of AMF. We address the importance of inoculum choice, plant choice, management practices and timing of inoculation for the successful manipulation of the resulting AMF community.

Comparative genomic and transcriptomic analyses reveal the hemibiotrophic stage shift of Colletotrichum fungi

Abstract: Summary Hemibiotrophic fungal plant pathogens represent a group of agronomically significant disease-causing agents that grow first on living tissue and then cause host death in later, necrotrophic growth. Among these, Colletotrichum spp. are devastating pathogens of many crops. Identifying expanded classes of genes in the genomes of phytopathogenic Colletotrichum, especially those associated with specific stages of hemibiotrophy, can provide insights on how these pathogens infect a large number of hosts. The genomes of Colletotrichum orbiculare, which infects cucurbits and Nicotiana benthamiana, and C. gloeosporioides, which infects a wide range of crops, were sequenced and analyzed, focusing on features with potential roles in pathogenicity. Regulation of C. orbiculare gene expression was investigated during infection of N. benthamiana using a custom microarray. Genes expanded in both genomes compared to other fungi included sequences encoding small, secreted proteins (SSPs), secondary metabolite synthesis genes, proteases and carbohydrate-degrading enzymes. Many SSP and secondary metabolite synthesis genes were upregulated during initial stages of host colonization, whereas the necrotrophic stage of growth is characterized by upregulation of sequences encoding degradative enzymes. Hemibiotrophy in C. orbiculare is characterized by distinct stage-specific gene expression profiles of expanded classes of potential pathogenicity genes.

Five phylogenetically close rice SWEET genes confer TAL effector‐mediated susceptibility to Xanthomonas oryzae pv. oryzae

Abstract: Bacterial plant-pathogenic Xanthomonas strains translocate transcription activator-like (TAL) effectors into plant cells to function as specific transcription factors. Only a few plant target genes of TAL effectors have been identified, so far. Three plant SWEET genes encoding putative sugar transporters are known to be induced by TAL effectors from rice-pathogenic Xanthomonas oryzae pv. oryzae (Xoo). We predict and validate that expression of OsSWEET14 is induced by a novel TAL effector, Tal5, from an African Xoo strain. Artificial TAL effectors (ArtTALs) were constructed to individually target 20 SWEET orthologs in rice. They were used as designer virulence factors to study which rice SWEET genes can support Xoo virulence. The Tal5 target box differs from those of the already known TAL effectors TalC, AvrXa7 and PthXo3, which also induce expression of OsSWEET14, suggesting evolutionary convergence on key targets. ArtTALs efficiently complemented an Xoo talC mutant, demonstrating that specific induction of OsSWEET14 is the key target of TalC. ArtTALs that specifically target individual members of the rice SWEET family revealed three known and two novel SWEET genes to support bacterial virulence. Our results demonstrate that five phylogenetically close SWEET proteins, which presumably act as sucrose transporters, can support Xoo virulence. © 2013 No claim to French Government works. © 2013 New Phytologist Trust.

The dual effect of abscisic acid on stomata

Abstract: The classical view that the drought-related hormone ABA simply acts locally at the guard cell level to induce stomatal closure is questioned by differences between isolated epidermis and intact leaves in stomatal response to several stimuli. We tested the hypothesis that ABA mediates, in addition to a local effect, a remote effect in planta by changing hydraulic regulation in the leaf upstream of the stomata. By gravimetry, porometry to water vapour and argon, and psychrometry, we investigated the effect of exogenous ABA on transpiration, stomatal conductance and leaf hydraulic conductance of mutants described as ABA-insensitive at the guard cell level. We show that foliar transpiration of several ABA-insensitive mutants decreases in response to ABA. We demonstrate that ABA decreases stomatal conductance and down-regulates leaf hydraulic conductance in both the wildtype Col-0 and the ABA-insensitive mutant ost2-2. We propose that ABA promotes stomatal closure in a dual way via its already known biochemical effect on guard cells and a novel, indirect hydraulic effect through a decrease in water permeability within leaf vascular tissues. Variability in sensitivity of leaf hydraulic conductance to ABA among species could provide a physiological basis to the isohydric or anisohydric behaviour.

Epigenetic variation creates potential for evolution of plant phenotypic plasticity

Abstract: Heritable variation in plant phenotypes, and thus potential for evolutionary change, can in principle not only be caused by variation in DNA sequence, but also by underlying epigenetic variation. However, the potential scope of such phenotypic effects and their evolutionary significance are largely unexplored. Here, we conducted a glasshouse experiment in which we tested the response of a large number of epigenetic recombinant inbred lines (epiRILs) of Arabidopsis thaliana--lines that are nearly isogenic but highly variable at the level of DNA methylation--to drought and increased nutrient conditions. We found significant heritable variation among epiRILs both in the means of several ecologically important plant traits and in their plasticities to drought and nutrients. Significant selection gradients, that is, fitness correlations, of several mean traits and plasticities suggest that selection could act on this epigenetically based phenotypic variation. Our study provides evidence that variation in DNA methylation can cause substantial heritable variation of ecologically important plant traits, including root allocation, drought tolerance and nutrient plasticity, and that rapid evolution based on epigenetic variation alone should thus be possible.

Drought predisposes piñon-juniper woodlands to insect attacks and mortality.

Abstract: Summary To test the hypothesis that drought predisposes trees to insect attacks, we quantified the effects of water availability on insect attacks, tree resistance mechanisms, and mortality of mature pinon pine (Pinus edulis) and one-seed juniper (Juniperus monosperma) using an experimental drought study in New Mexico, USA. The study had four replicated treatments (40 × 40 m plot/replicate): removal of 45% of ambient annual precipitation (H2O−); irrigation to produce 125% of ambient annual precipitation (H2O+); a drought control (C) to quantify the impact of the drought infrastructure; and ambient precipitation (A). Pinon began dying 1 yr after drought initiation, with higher mortality in the H2O− treatment relative to other treatments. Beetles (bark/twig) were present in 92% of dead trees. Resin duct density and area were more strongly affected by treatments and more strongly associated with pinon mortality than direct measurements of resin flow. For juniper, treatments had no effect on insect resistance or attacks, but needle browning was highest in the H2O− treatment. Our results provide strong evidence that ≥ 1 yr of severe drought predisposes pinon to insect attacks and increases mortality, whereas 3 yr of the same drought causes partial canopy loss in juniper.

Confronting model predictions of carbon fluxes with measurements of Amazon forests subjected to experimental drought

Abstract: Considerable uncertainty surrounds the fate of Amazon rainforests in response to climate change. Here, carbon (C) flux predictions of five terrestrial biosphere models (Community Land Model version 3.5 (CLM3.5), Ecosystem Demography model version 2.1 (ED2), Integrated BIosphere Simulator version 2.6.4 (IBIS), Joint UK Land Environment Simulator version 2.1 (JULES) and Simple Biosphere model version 3 (SiB3)) and a hydrodynamic terrestrial ecosystem model (the Soil-Plant-Atmosphere (SPA) model) were evaluated against measurements from two large-scale Amazon drought experiments. Model predictions agreed with the observed C fluxes in the control plots of both experiments, but poorly replicated the responses to the drought treatments. Most notably, with the exception of ED2, the models predicted negligible reductions in aboveground biomass in response to the drought treatments, which was in contrast to an observed c. 20% reduction at both sites. For ED2, the timing of the decline in aboveground biomass was accurate, but the magnitude was too high for one site and too low for the other. Three key findings indicate critical areas for future research and model development. First, the models predicted declines in autotrophic respiration under prolonged drought in contrast to measured increases at one of the sites. Secondly, models lacking a phenological response to drought introduced bias in the sensitivity of canopy productivity and respiration to drought. Thirdly, the phenomenological water-stress functions used by the terrestrial biosphere models to represent the effects of soil moisture on stomatal conductance yielded unrealistic diurnal and seasonal responses to drought.

Increased vapor pressure deficit due to higher temperature leads to greater transpiration and faster mortality during drought for tree seedlings common to the forest-grassland ecotone.

Abstract: Summary Tree species growing along the forest–grassland ecotone are near the moisture limit of their range. Small increases in temperature can increase vapor pressure deficit (VPD) which may increase tree water use and potentially hasten mortality during severe drought. We tested a 40% increase in VPD due to an increase in growing temperature from 30 to 33°C (constant dewpoint 21°C) on seedlings of 10 tree species common to the forest–grassland ecotone in the southern Great Plains, USA. Measurement at 33 vs 30°C during reciprocal leaf gas exchange measurements, that is, measurement of all seedlings at both growing temperatures, increased transpiration for seedlings grown at 30°C by 40% and 20% for seedlings grown at 33°C. Higher initial transpiration of seedlings in the 33°C growing temperature treatment resulted in more negative xylem water potentials and fewer days until transpiration decreased after watering was withheld. The seedlings grown at 33°C died 13% (average 2 d) sooner than seedlings grown at 30°C during terminal drought. If temperature and severity of droughts increase in the future, the forest–grassland ecotone could shift because low seedling survival rate may not sufficiently support forest regeneration and migration.

Nematode effector proteins: an emerging paradigm of parasitism

Abstract: Phytonematodes use a stylet and secreted effectors to modify host cells and ingest nutrients to support their growth and development. The molecular function of nematode effectors is currently the subject of intense investigation. In this review, we summarize our current understanding of nematode effectors, with a particular focus on proteinaceous stylet-secreted effectors of sedentary endoparasitic phytonematodes, for which a wealth of information has surfaced in the past 10 yr. We provide an update on the effector repertoires of several of the most economically important genera of phytonematodes and discuss current approaches to dissecting their function. Lastly, we highlight the latest breakthroughs in effector discovery that promise to shed new light on effector diversity and function across the phylum Nematoda.

Integrative invasion science: model systems, multi-site studies, focused meta-analysis and invasion syndromes.

Abstract: Summary Invasion scienceis averyactive subdisciplineofecology. However,somescientists contendthat theoretical integration has been limited and that predictive power remains weak. This paper, focusing on plants, proposes a new multi-pronged research strategy that builds on recent advances in invasion science. More intensive studies on particular model organisms and ecosystems are needed to improve our understanding of the full suite of interacting factors that influence invasions ('model system research'). At the same time, comparative studies across many study systems are essential for unravelling the context-dependencies of insights that emerge from particular studies ('multi-site studies'); and quantitative synthesis based on large datasets should be constrained to well-defined theoretical domains ('focused meta-analysis'). Wealsosuggestwaysforbetterintegrationofinformationaboutspeciesbiologyandecosystem characteristics('invasionsyndromes').Weexpectthataresultingtheoryofinvasionswillneedto beconceivedasasomewhatheterogeneousconglomerateofelementsofvaryinggeneralityand predictive power: laws that apply to well-specified domains, general concepts and theoretical frameworks that can guide thinking in research and management, and in-depth knowledge about the drivers of particular invasions.

Thirst beats hunger – declining hydration during drought prevents carbon starvation in Norway spruce saplings

Abstract: Summary � Drought-induced tree mortality results from an interaction of several mechanisms. Plant water and carbon relations are interdependent and assessments of their individual contributions are difficult. Because drought always affects both plant hydration and carbon assimilation, it is challenging to disentangle their concomitant effects on carbon balance and carbon translocation. Here, we report results of a manipulation experiment specifically designed to separate drought effects on carbon and water relations from those on carbon translocation. � In a glasshouse experiment, we manipulated the carbon balance of Norway spruce saplings exposed to either drought or carbon starvation (CO2 withdrawal), or both treatments, and compared the dynamics of carbon exchange, allocation and storage in different tissues. � Drought killed trees much faster than did carbon starvation. Storage C pools were not depleted at death for droughted trees as they were for starved, well-watered trees. Hence drought has a significant detrimental effect on a plant’s ability to utilize stored carbon. � Unless they can be transported to where they are needed, sufficient carbon reserves alone will not assure survival of a drought except under specific conditions, such as moderate drought, or in species that maintain plant water relations required for carbon re-mobilization.