Academic literature on the topic 'Plant adaptation'

Plants in their natural habitats adapt to drought stress in the environment through a variety of mechanisms, ranging from transient responses to low soil moisture to major survival mechanisms of escape by early flowering in absence of seasonal rainfall. However, crop plants selected by humans to yield products such as grain, vegetable, or fruit in favorable environments with high inputs of water and fertilizer are expected to yield an economic product in response to inputs. Crop plants selected for their economic yield need to survive drought stress through mechanisms that maintain crop yield. Studies on model plants for their survival under stress do not, therefore, always translate to yield of crop plants under stress, and different aspects of drought stress response need to be emphasized. The crop plant model rice (Oryza sativa) is used here as an example to highlight mechanisms and genes for adaptation of crop plants to drought stress.

In this short review, I will first summarize criteria by which environments can be considered “cold”, with plant stature (size, height above ground) playing a central role for the climate actually experienced. Plants adapted to such environments have to cope with both extremes and with gradual influences of low temperature. The first requires freezing resistance, which is tightly coupled to developmental state (phenology) and prehistory (acclimation). Gradual low temperature constraints affect the growth process (meristems) long before they affect photosynthetic carbon gain. Hence, plants growing in cold climates are commonly not carbon limited.

Winter survival is a complex trait that does not solely rely on the plant's ability to withstand the direct effects of extreme cold temperatures. During long overwintering periods, plants are exposed to multiple abiotic (ice encasement, frost heave, desiccation, anoxia) and biotic (snow mould and other psychrophylic pathogens) stresses. Tolerance to these various stresses is based in part on shared adaptive traits and, consequently, cross-adaptation to environmental stresses is a key aspect of plant adaptation to cold. Increasing evidence of multiple functions for stress-induced proteins in overwintering plants confirms the need for a global approach in the analysis of adaptive mechanisms. From that perspective, the valorization of rapidly increasing knowledge on the molecular and genetic basis of plant and microbe adaptations to cold will demand multidisciplinary collaborations. Climate change will also need to be taken into account to identify the adaptive traits that will be required for agricultural and forest plants to survive winter in the future. More studies at the global and regional scales will be needed to assess the potential impact of climate warming on plant adaptation to winter and their interactions with low-temperature pathogens.Key words: cold adaptation, psychrophylic microorganisms, climate change, fall dormancy, low-temperature plant–microbe interactions, cold-adaptation genomics.

Understanding how local adaptation arises within species is essential when making predictions about how populations may respond to environmental change. Serpentine soils present an ideal system to study the processes leading to local adaptation in plant populations. Theses soils have a scattered distribution, are deficient in several nutrient minerals, have high heavy metal content, and a poor water holding capacity; all factors contributing to a strong selective pressure against plant growth. This thesis investigates local adaptation to serpentine soil in populations of Arabidopsis lyrata ssp. petraea, a perennial of low competition habitats and which within Scotland occurs both on and off serpentine soil. Trait measures were made of plants from twelve populations across the species’ Scottish range to explore if there are trait patterns which are particular to the serpentine sites, finding that climatic and soil factors both contributed to observed trait patterns. In a controlled growth room experiment the response of five of the populations to growth on serpentine and non-serpentine soil treatments was assessed to determine if trait differences were the result of local adaptation. Plants from serpentine populations survived and performed well on all treatments, while non-serpentine plants showed reduced survival and poor growth on serpentine soil. Combined with patterns of differentiation in microsatellite markers indicating that there is not a single serpentine genetic strain of Scottish A. l. petraea it appears that serpentine populations have adapted to serpentine soil, and separately more than once. Finally, an individual based computer model was developed to explore the evolutionary consequences of local adaptation in a changing environment, finding that the strength of local adaptation, gene flow, and population size are all important determinants of population survival. These results have implications for conservation management practices used to protect small populations in changing environments; especially where local adaption is likely.

Plant morphology and function are sensitive to rising atmospheric carbon dioxide (CO2) concentrations, but evidence that CO2 concentration can act as a selective pressure driving evolution is sparse. Plants originating from naturally high CO2 springs are subjected to elevated CO2 concentration over multiple generations, providing an opportunity to predict how adaptation to future atmospheres may occur, with important implications for future plant conservation and crop breeding strategies. Using Plantago lanceolata L. from such a site (the ‘spring’ site) and from an adjacent ambient CO2 site (‘control’ site), and growing the populations in ambient and elevated CO2 at 700 μmol mol-1, I have characterised, for the first time, the functional and population genomics, alongside morphology and physiology, of plant adaptation to elevated CO2 concentrations. Growing plants in elevated CO2 caused relatively modest changes in gene expression, with fewer changes evident in the spring than control plants (33 vs 131 genes differentially expressed [DE], in spring and control plants respectively). In contrast, when comparisons were made between control and spring plants grown in either ambient or elevated CO2, there were a much larger number of loci showing DE (689 in the ambient and 853 in the elevated CO2 environment). Population genomic analysis revealed that genetic differentiation between the spring and control plants was close to zero with no fixed differences, suggesting that plants are adapted to their native CO2 environment at the level of gene expression. Growth at elevated CO2 led to an unusual phenotype, with an increase in stomatal density and index in the spring, but not in control plants. Focussing on previously characterised stomatal patterning genes revealed significant DE (FDR < 0.05) between spring and control plants for three loci (YODA, CDKB1;1, and SCRM2) and between ambient and elevated CO2 for four (ER, YODA, MYB88, and BCA1). We propose that the up-regulation in spring plants of two positive regulators of stomatal numbers (SCRM2 and CDKB1;1) act here as key controllers of stomatal adaptation to elevated CO2 on an evolutionary timescale. Significant transcriptome reprogramming of the photosynthetic pathway was identified, with an overall decrease in expression across the pathway in control plants, and an increase in spring plants, in response to elevated CO2. This was followed up by physiological measurements, where a significant increase (P < 0.05) in photosynthetic capacity and regeneration rate was exhibited in spring plants, compared to control plants, at both elevated and ambient CO2 concentrations. Through this comprehensive analysis, we have identified the basis of plant adaptation to elevated CO2 likely to occur in the future.

This study lends strong support to the idea that members of Bromeliaceae have undergone a recent adaptive radiation, and therefore show that, at least in part, diversity in the tropics is due to a fast speciation rate and that the tropics can be a "cradle" for new diversification and exploitation of varying ecological niches through the diversification of ecophysiological traits within a lineage.

The obligately outbreeding root hemiparasite Striga hermonthica (Orobanchaceae)is a serious threat to subsistence agriculture in sub-Saharan Africa. Resistance to this parasite in its crop hosts, such as rice, sorghum and maize, is not common, and the evolution of host adaptations that are able to overcome new sources of resistance is an ever-present risk. Research into host adaptation in S. hermonthica has generally sought to correlate the genetic relationship between Striga individuals with host identity; however, such approaches must be supported by lab-based evidence of host adaptation, otherwise ad hoc field sampling may result in the confounding of host identity with isolation-by-distance. Additionally, genetic variation used to reconstruct relationships is unlikely to provide an insight into relationships at functional loci underlying host adaptation. In this thesis, I use a range of new approaches to investigate several different aspects of parasite adaptation in the S. hermonthica-Sorghum bicolor pathosystem. Host adaptations, or pre-adaptations, are commonly revealed using tests for differential virulence between Striga populations and host genotypes; that is, by demonstrating population-level genotype-by-genotype interactions. Evidence for such interactions was found between three West African populations of S. hermonthica and five sorghum cultivars. These interactions were shown to be strongest at the parasite post-attachment life stage, and to depend on the parasite virulence metric used. Environmental influences on host-parasite interactions were strong and variable between years and sites. Candidate genetic loci for virulence, responding to selection in a micro-evolutionary fashion, were identified by Fst differentiation-based approaches ('outlier analyses') that aim to uncover associations between particular loci and environmental drivers, such as host identity. An AFLP outlier analysis was used on Striga plants parasitising nine sorghum cultivars in a field trial in Burkina Faso. Significant locus-specific differentiation was detected at 14 out of 1275 loci. However, predicted allele frequencies at these loci did not correlate with a field measure of Striga virulence across host-selected sub-populations. Simulation results suggested that the estimated levels of Fst at outlying loci could mean that alleles underlying host adaptation exist at intermediate frequencies in populations. A three-generation pedigree, created from a cross between S. hermonthica individuals from an East African and a West African population, enabled further insights into the genetics of adaptation. Individuals from a pseudo-backcrossed F1 (BCF1) generation, grown on two different sorghum hosts and in axenic culture, indicated significant host-related segregation distortion. Analyses of virulence in the BCF1 also provided strong evidence for epistasis, and for an effect of maternal identity. A second outlier analysis of host adaptation, investigating the East African population used in the pedigree, indicated some correspondence between outlier loci and loci found to be differentially segregating between different hosts in the BCF1 generation, and demonstrated the differing genomic extents of these phenomena. The results accumulated across these experiments provide evidence for a complex, polygenic basis to virulence in S. hermonthica.

Multi environment trials conducted over mapping population are often used to test genotypes in a set of environments that represent the target environmental range. The first part of this work is the evaluation of the ‘Nure’ x ‘Tremois’ double-­‐haploid mapping population, together with an association panel comprising 185 barley varieties representative of the barley germplasm cultivated in the Mediterranean basin. Plant material was tested across eighteen site by year field trials combination, in six countries across the Mediterranean basin. Trials were growth at sites contrasting for natural rainfall (high vs low on the base of past meteorological data) or at the same site with one being rainfed and the other with supplementary irrigation. Trials conducted for two years in each one of the sites and this allowed tocollect a huge data series comprising agronomical traits defining grain yield and yield components, phenological and environmental data, subsequently used to identify genomic regions involved in barley adaptation. The 118 doubled haploid lines of the mapping population were genotyped with Diversity Array Technology® (DaRT) marker assay and subsequently a total of 15 CAPS and SSCP marker for candidate genes involved in phenology regulation and abiotic stress response were added to the linkage map based on DaRT markers. Data collected were firstly used to perform QTLs analysis with composite interval mapping for any environment/ trait combination, results showed eight QTLs for grain yield, days to heading and grain yield components. . The two mostly frequents QTLs for grain yield and days to heading were located on barley chromosome 1H (3 trials), 2H (8 trials) and 5H (5 trials) overlapping respectively HvFT3 gene, the earliness per se locus (eam6/Eps-­‐2) and the vernalization gene Vrn_H1. A further QTL multi-­‐environment analysis was performed and revealed that across the 18 field trials QTL for eam6/Eps-­‐2 (2H) and Vrn-­‐H1 (5H) were commons for days to heading and grain yield. We use all the environmental information collected to check QTLs sensitivities to co-­‐environmental co-­‐variables. Most of significant associations collected were related to temperature and temperature-­‐based variables troughtout the growing cycle. Eam6/Eps-­‐2 showed non-­‐crossover QTL.E interaction, while for Vrn-­‐H1 crossover interactions were revealed. The 185 barley accession were genotyped with 1536 SNPs and data collected for this population for cold resistance in two field trials in Spain an Italy, the first trial was characterized by an exceptional winter, while the second was previously know has frost-­‐prone environment. Results from genome wide association analysis showed 13 positive associations with specific genomic regions. Interestingly several of these QTL were coincident with the position of previously mapped loci for cold tolerance, on chromosomes 2HL, 4HL and 5HL.
Els assajos en localitats múltiplas de poblacions de mapeo s'utilitzen freqüentment per a testar genotips en un conjunt d'ambients representatius de la condicions climàtiques on es volen introduir aquests genotips. La primera part d'això treball ha estat l'avaluació de la població de mapeo ‘Nure x Tremois’ constituïda de 118 de doble haploides d'ordi, juntament amb panell d'associació que comprèn 185 varietats d'ordi representatives del germoplasma conreat en la conca Mediterrània. El material vegetal ha estat assajat en una combinació de divuit camps per any desllorigats en sis països de la conca mediterrània. Els assajos s'han portat a terme en camps amb diferent disponibilitat d'aigua, classificats sobre la base de les dades relatives a les freqüència i quantitat de les precipitacions o en el mateix lloc amb un camp en secà i altre regat. Els assajos es van portar a terme per dos anys en cada localitat i això va permetre la recollida d'un gran volum de dades que comprenen caràcters agronómicos relacionats amb rendiment i components del rendiment, dades fenológicos i ambientals. Aquestes dades es van utilitzar després per a la identificació de regions genomicas involucrades en l'adaptació de l'ordi a l'ambient. Els 118 dobles haploides de la població ‘Nure x Tremois’ es genotiparon amb marcadors DaRT (Diversity Array Technology), després un set de 15 marcadors CAPS I SCCP per a gens candidats involucrats en la regulació de les fases fenológicas van ser afegits al mapa de lligament construït amb els marcadors DaRT. Les dades van ser utilitzats per a fer una anàlisi de QTL amb procediment ‘Composite Interval Mapping’ para cada combinació ambienti/ caràcter. Es van trobar diversos QTLs per rendiment i data d'espigolat i components del rendiment. Els QTL mes freqüents trobats per rendiment i data de floració i components del rendiment estan localitzats en els cromosomes 1H (3 camps), 2H (8 camps) i 5H (5 camps) coincidents respectivament amb HvFT3 locus, eam6/Eps-­‐2 (earliness per se) locus i amb el locus de vernalización Vrn-­‐H1. Una ulterior anàlisi de QTL feta amb el mètode “Multi Environment Trial” ha revelat que els QTL localitzats en el locus eam6/Eps-­‐2 (cromosoma 2H) i Vrn-­‐H1 (cromosoma 5H) són comunes per rendiment i data de floració en els 18 camps d'assaig. Per això utilitzem tots el dades ambientals col·leccionades durant tot el cicle del cultiu per a investigar la sensibilitat de dites QTL a les co-­‐variables ambientals. La majoria de les associacions oposades estan relacionades amb temperatures i variables relacionades amb aquestes. Eam6/Eps-­‐2 mostra una interacció de tipus quantitatiu amb aquestes variables mentre Vrn-­‐H1 mostra una interacció de tipus qualitatiu amb aquestes variables. Les 185 varietats assajades van ser genotipadas amb 185 SNPs i fenotipadas per resistència a fred en dos assajos uneixo a Espanya i altre a Itàlia. El primer assaig va ser caracteritzat per un hivern excepcionalment fred, mentre el d'Itàlia ha estat utilitzat en passat per testar resistència a fred a causa de els hiverns rígids que solen registrar-­‐se en aquesta localitat. Les dades van ser utilitzats per a portar a terme la analisis GWAS “Genome Wide Association Analysis” . Els resultats van permetre identificar 13 regions genomicas involucrades en la resistència a frio. Entre elles tres regions coincideixen amb loci ja mapeados i coneguts per ser involucrats en la resposta a frio en los cromosomes 2HL, 4HL i 5HL.
Los ensayos en localidades múltiplas de poblaciones de mapeo se utilizan frecuentemente para testar genotipos en un conjunto de ambientes representativos de la condiciones climáticas donde se quieren introducir dichos genotipos. La primera parte de esto trabajo ha sido la evaluación de la población de mapeo ‘Nure x Tremois’ constituida de 118 de doble haploides de cebada, junto con panel de asociación que comprende 185 variedades de cebada representativas del germoplasma cultivado en la cuenca Mediterránea. El material vegetal ha sido ensayado en una combinación de dieciocho campos por año dislocados en seis países de la cuenca mediterránea. Los ensayos se han llevado a cabo en campos con diferente disponibilidad de agua, clasificados en base a los datos relativos a las frecuencia y cantidad de las precipitaciones o en el mismo sitio con un campo en secano y otro regado. Los ensayos se llevaron a cabo por dos años en cada localidad y esto permitió la recogida de un gran volumen de datos que comprenden caracteres agronómicos relacionados con rendimiento y componentes del rendimiento, datos fenológicos y ambientales. Dichos datos se utilizaron después para la identificación de regiones genomicas involucradas en la adaptación de la cebada al ambiente. Los 118 dobles haploides de la población ‘Nure x Tremois’ se genotiparon con marcadores DaRT (Diversity Array Technology), después un set de 15 marcadores CAPS Y SCCP para genes candidatos involucrados en la regulación de las fases fenológicas fueron añadidos al mapa de ligamento construido con los marcadores DaRT. Los datos fueron utilizados para hacer una análisis de QTL con procedimiento ‘Composite Interval Mapping’ para cada combinación ambiente/ carácter. Se encontraron varios QTLs por rendimiento y fecha de espigado y componentes del rendimiento. Los QTL mas frecuentes encontrados por rendimiento y fecha de floración y componentes del rendimiento están localizados en los cromosomas 1H (3 campos), 2H (8 campos) y 5H(5 campos) coincidentes respectivamente con HvFT3 locus, eam6/Eps-­‐2 (earliness per se) locus y con el locus de vernalización Vrn-­‐H1. Una ulterior análisis de QTL hecha con el método “Multi Environment Trial” ha revelado que los QTL localizados en el locus eam6/Eps-­‐2 (cromosoma 2H) y Vrn-­‐H1 (cromosoma 5H) son comunes por rendimiento y fecha de floración en los 18 campos de ensayo. Por esto utilizamos todos lo datos ambientales coleccionadas durante todo el ciclo del cultivo para investigar la sensibilidad de dichos QTL a las co-­‐variables ambientales. La mayoría de las asociaciones encontradas están relacionadas con temperaturas y variables relacionadas con estas. Eam6/Eps-­‐2 muestra una interacción de tipo cuantitativo con dichas variables mientras Vrn-­‐H1 muestra una interacción de tipo cualitativo con dichas variables. Las 185 variedades ensayadas fueron genotipadas con 185 SNPs y fenotipadas por resistencia a frío en dos ensayos uno en España y otro en Italia. El primer ensayo fue caracterizado por un invierno excepcionalmente frío, mientras el de Italia ha sido utilizado en pasado por testar resistencia a frío debido a los inviernos rígidos que suelen registrarse en dicha localidad. Los datos fueron utilizados para llevar a cabo la analisis GWAS “Genome Wide Association Analysis”. Los resultados permitieron identificar 13 regiones genomicas involucradas en la resistencia a frio. Entre ellas tres regiones coinciden con loci ya mapeados y conocidos por ser involucrados en la respuesta a frio en los cromosomas 2HL, 4HL y 5HL.

Species extinction rates are causing alarm. Anthropogenic distortion of the climate system is rapidly altering the natural environment. Arabidopsis thaliana is a model species in molecular biology with widespread wild populations showing functional diversity however its ecology and evolution is poorly understood. Faced with a changing natural world, what is the adaptive potential of the model plant species Arabidopsis thaliana? This thesis focuses on the interactions of genotypes, phenotypes and environments to assess the current state of adaptation in this vagile species and to identify mechanisms for rapid adaptation to future stress, focusing on plant pathogens. Here I show that A. thaliana populations in England exhibit evidence of local adaptation and genetic structure. A large common garden experiment using genotypes gathered in natural habitats revealed functional fitness differences in genotype-by-environment interactions. Wild populations showed differential representation of RPM1 alleles suggesting non-random processes are responsible for the exhibited patterns. A further common garden experiment demonstrated ‘home site advantage’ through a correlation between fitness and home site climate, which suggests that local adaptation had occurred. Phenotypic plasticity and mechanisms for rapid adaptation could be essential for plant survival under predicted climate change. Using Xanthomonas spp. as xenopathogens, I show differing levels of pre-adaptation for pathogen response exists in wild UK populations of A. thaliana. By using a multi-generation study, I found some evidence that epigenetic modification enabled rapid adaptation to pathogen stress. Finally, I compared the metabolic expressions of phenotype among genotypes in two artificial environments. Environmental effects detected by this method are far greater than genetic ones, suggesting that metabolic plasticity can underpin environmental adaptation. Taken together, my results suggest that wild populations of A. thaliana contain a range of mechanisms for rapid adaptation to environmental change. If these capacities are general, my work offers a note of optimism about the fate of some wild plant species in the face of global climate change. Additionally, as A. thaliana is a model species in genomics, my findings may facilitate future exploitation of these traits by crop geneticists.

AbstractThe vulnerability of agriculture systems in Africa to climate change is directly and indirectly affecting the availability and diversity of plants and plant products available in local markets. In this chapter, markets in Benin City and environs were assessed to document the availability of plants and plant products. Markets were grouped into urban, suburban, and rural with each group having four markets. Majority of the plant and plant product vendors were women and 88 plant species belonging to 42 families were found. Their scientific and common names were documented as well as the parts of the plant and associated products available in the markets. Most of the plant and plant products found in local markets belong to major plant families. Urban markets had the highest diversity of plants and plant products. Three categories of plants and plant products were documented. Around 67% of the plants and plant products were categorized as whole plant/plant parts, 28% as processed plant parts, while 5% as reprocessed plant/plant parts. It was revealed that 86% of these plants are used as foods, 11% are for medicinal purposes, while 3% is used for other purposes. About 35% of plants and plant products across the markets were fruits, which is an indication that city and environs are a rich source of fruits. The local knowledge and practices associated with the plants and plant products can contribute towards formulating a strategic response for climate change impacts on agriculture, gender, poverty, food security, and plant diversity.

The AP1000® plant is an 1100-MWe pressurized water reactor (PWR) with passive safety features and extensive plant simplifications that enhance construction, operation, maintenance and safety. One of the key design approaches in the AP1000 plant is to use passive features to mitigate design basis accidents. Active defense-in-depth (DiD) features provide investment protection, reduce the demands on the passive features and support the PRA. The passive features are classified as safety-related in the US. The active defense-in-depth features are classified as non-safety (with supplemental requirements) in the US. The AP1000 design has incorporated a standardization approach, which together with the level of safety achieved by the passive safety features, results in a plant design that can be applied to different geographical regions with varying regulatory standards and utility expectations without major changes. While the first deployments of the AP1000 plant are ongoing in China and the United States, Westinghouse has remained active in also pursuing European opportunities for the AP1000 plant. In particular, Westinghouse has cooperated for almost two decades with European utilities to ensure adaptation of the AP1000 plant to the European market. This cooperation has resulted in progress towards AP1000 plant deployment in European countries. The AP1000 plant is recognized worldwide and has been reviewed by regulators around the world, including China, the United Kingdom (UK), Canada as well as the US. The AP1000 PWR is the only Generation III+ reactor design to obtain final design approval from the United States Nuclear Regulatory Commission (US NRC) and interim approval from UK regulatory authorities as part of the Generic Design Assessment (GDA) process. It is the only technology to be licensed for construction in the United States in more than 30 years, and the only Generation III+ technology worldwide to receive an operating license, as well as construction approval in China. The AP1000 plant has been independently assessed and confirmed to meet the requirements of the European Utilities Requirements (EUR) document and the Electric Power Research Institute (EPRI) Advanced Light Water Reactor Utility Requirements Document (URD). The AP1000 plant has also been successfully assessed against multiple European industry guidelines such as the WENRA safety objectives, the IAEA safety standards, the ENSREG stress tests and the UK Weightman Report. In support of multiple ongoing request for proposal (RFP) and pre-RFP activities in European countries, Westinghouse has focused design effort and customer interactions in several European countries to adapt the AP1000 plant to European requirements. Review of the AP1000 plant design with regulators around the world, European Standards compliance activities, and continued cooperation and interaction with European Utilities provide confidence that the AP1000 plant can be successfully licensed and deployed in Europe. The AP1000 50Hz standard plant design (also referred to as European Passive Standard or EPS) is the resulting adaptation of the AP1000 60 Hz US standard plant design to European market needs and requirements, addressing both customer input from such programs as the European Passive Plant (EPP) program in addition to regulatory and Utility needs identified though RFP and pre-RFP activities. The AP1000 50Hz standard plant design retains the overall AP1000 plant design (safe, simple, standard), the use of proven components and its cost, safety and operability advantages, while incorporating some changes to adapt to the European environment. This paper will discuss some of the key changes that have been incorporated into the AP1000 50Hz plant design as necessary to adapt to the European market and demonstrate that the vast majority of the standard AP1000 plant design being built in China and the US is not impacted.

Original Objectives: 1. Purify and biochemically characterize RB60 orthologs in higher plant chloroplasts; 2. Clone the gene(s) encoding plant RB60 orthologs and determine their structure and expression; 3. Manipulate the expression of RB60; 4. Assay the effects of altered RB60 expression on thylakoid biogenesis and photosynthetic function in plants exposed to different light conditions. In addition, we also examined the gene structure and expression of RB60 orthologs in the non-vascular plant, Physcomitrella patens and cloned the poly(A)-binding protein orthologue (43 kDa RB47-like protein). This protein is believed to a partner that interacts with RB60 to bind to the psbA5' UTR. Thus, to obtain a comprehensive view of RB60 function requires analysis of its biochemical partners such as RB43. Background & Achievements: High levels of sunlight reduce photosynthesis in plants by damaging the photo system II reaction center (PSII) subunits, such as D1 (encoded by the chloroplast tpsbAgene). When the rate of D1 synthesis is less than the rate of photo damage, photo inhibition occurs and plant growth is decreased. Plants use light-activated translation and enhanced psbAmRNA stability to maintain D1 synthesis and replace the photo damaged 01. Despite the importance to photosynthetic capacity, these mechanisms are poorly understood in plants. One intriguing model derived from the algal chloroplast system, Chlamydomonas, implicates the role of three proteins (RB60, RB47, RB38) that bind to the psbAmRNA 5' untranslated leader (5' UTR) in the light to activate translation or enhance mRNA stability. RB60 is the key enzyme, protein D1sulfide isomerase (Pill), that regulates the psbA-RN :Binding proteins (RB's) by way of light-mediated redox potentials generated by the photosystems. However, proteins with these functions have not been described from higher plants. We provided compelling evidence for the existence of RB60, RB47 and RB38 orthologs in the vascular plant, Arabidopsis. Using gel mobility shift, Rnase protection and UV-crosslinking assays, we have shown that a dithiol redox mechanism which resembles a Pill (RB60) activity regulates the interaction of 43- and 30-kDa proteins with a thermolabile stem-loop in the 5' UTR of the psbAmRNA from Arabidopsis. We discovered, in Arabidopsis, the PD1 gene family consists of II members that differ in polypeptide length from 361 to 566 amino acids, presence of signal peptides, KDEL motifs, and the number and positions of thioredoxin domains. PD1's catalyze the reversible formation an disomerization of disulfide bonds necessary for the proper folding, assembly, activity, and secretion of numerous enzymes and structural proteins. PD1's have also evolved novel cellular redox functions, as single enzymes and as subunits of protein complexes in organelles. We provide evidence that at least one Pill is localized to the chloroplast. We have used PDI-specific polyclonal and monoclonal antisera to characterize the PD1 (55 kDa) in the chloroplast that is unevenly distributed between the stroma and pellet (containing membranes, DNA, polysomes, starch), being three-fold more abundant in the pellet phase. PD1-55 levels increase with light intensity and it assembles into a high molecular weight complex of ~230 kDa as determined on native blue gels. In vitro translation of all 11 different Pill's followed by microsomal membrane processing reactions were used to differentiate among PD1's localized in the endoplasmic reticulum or other organelles. These results will provide.1e insights into redox regulatory mechanisms involved in adaptation of the photosynthetic apparatus to light stress. Elucidating the genetic mechanisms and factors regulating chloroplast photosynthetic genes is important for developing strategies to improve photosynthetic efficiency, crop productivity and adaptation to high light environments.

Annual plants have developed a range of different mechanisms to avoid flowering (exposure of reproductive organs to the environment) under adverse environmental conditions. Seasonal environmental events such as gradual changes in day length and temperature affect the timing of transition to flowering in many annual and perennial plants. Research in Arabidopsis and additional species suggest that some environmental signals converge on transcriptional regulation of common floral integrators such as FLOWERING LOCUS T (FT). Here we studied environmental induction of flowering in the model legume Medicago truncatula. Similarly to Arabidopsis, the transition to flowering in M. truncatula is hastened by long photoperiods and long periods of vernalization (4°C for 2-3 weeks). Ecotypes collected in Israel retain a vernalization response even though winter temperatures are way above 4°C. Here we show that this species is also highly responsive (flowers earlier) to mild ambient temperatures up to 19°C simulating winter conditions in its natural habitat. Physiological experiments allowed us to time the transition to flowering due to low temperatures, and to compare it to vernalization. We have made use of natural variation, and induced mutants to identify key genes involved in this process, and we provide here data suggesting that an FT gene in M.truncatula is transcriptionally regulated by different environmental cues. Flowering time was found to be correlated with MtFTA and MtFTB expression levels. Mutation in the MtFTA gene showed a late flowering phenotype, while over-expressing MtFTA in Arabidopsis complemented the ft- phenotype. We found that combination of 4°C and 12°C resulted in a synergistic increase in MtFTB expression, while combining 4°C and long photoperiods caused a synergistic increase in MtFTA expression. These results suggest that the two vernalization temperatures work through distinct mechanisms. The early flowering kalil mutant expressed higher levels of MtFTA and not MtFTB suggesting that the KALIL protein represses MtFTA specifically. The desert ecotype Sde Boker flowers earlier in response to short treatments of 8-12oc vernalization and expresses higher levels of MtFTA. This suggests a possible mechanism this desert ecotype developed to flower as fast as possible and finish its growth cycle before the dry period. MtFTA and FT expression are induced by common environmental cues in each species, and expression is repressed under short days. Replacing FT with the MtFTA gene (including regulatory elements) caused high MtFTA expression and early flowering under short days suggesting that the mechanism used to repress flowering under short days has diversified between the two species.The circadian regulated gene, GIGANTEA (GI) encodes a unique protein in Arabidopsis that is involved in flowering mechanism. In this research we characterized how the expression of the M.truncatula GI ortholog is regulated by light and temperature in comparison to its regulation in Arabidopsis. In Arabidopsis GI was found to be involved in temperature compensation to the clock. In addition, GI was found to be involved in mediating the effect of temperature on flowering time. We tested the influence of cold temperature on the MtGI gene in M.truncatula and found correlation between MtGI levels and extended periods of 12°C treatment. MtGI elevation that was found mostly after plants were removed from the cold influence preceded the induction of MtFT expression. This data suggests that MtGI might be involved in 12°C cold perception with respect to flowering in M.truncatula. GI seems to integrate diverse environmental inputs and translates them to the proper physiological and developmental outputs, acting through several different pathways. These research enabled to correlate between temperature and circadian clock in M.truncatula and achieved a better understanding of the flowering mechanism of this species.

In order to advance our understanding towards potential biotechnology improvement of plant performance, we studied root responses to limited P in two different plants, Arabidopsis and tomato. Arabidopsis is among the most studied model plants that allows rapid application of molecular and developmental experiments while tomato is an important crop, with application in agriculture. Using Arabidopsis we found that steroid hormones modulate the extent of root elongation in response to limited P, by controlling the accumulation of iron in the root. We also found that the availability of P and iron control the activity of the steroid hormone in the root. Finally, we revealed the genes involved in this nutrient-hormone interaction. Hence, the ferroxidase LPR1 that promotes iron accumulation in response to low P is repressed by the transcription factor BES1/BZR1. Low P inhibits the steroid hormone pathway by enhancing the accumulation of BKI1. High levels of BKI1 inhibit the activity of the steroid hormone receptor at the cell surface and iron accumulation increases inside the root, resulting in a slow growth. Together, the extent of root elongation depends on interactions between an internal cue (steroid hormone) and cues derived from the availability of P and iron in the environment. Using tomato, we found that the response of two cultivated tomato varieties (M82 and New Yorker) to limited P is distinct from that of the wild species, Solanumpennellii. This is implicated at both the levels of root development and whole plant physiology. Specifically, while the root system architecture of cultivated tomato is modulated by limited P availability, that of the wild type species remained unaffected. The wild species appears to be always behaving as if it is always in phosphate deprived conditions, despite sufficient levels of phosphate. Hyper-accumulation of metals appears to mediate this response. Together, this knowledge will be used to isolate new genes controlling plant adaptation to limited P environment.

Original objectives: [a] Screen an array of chickpea and wild annual Cicer germplasm for winter survival. [b] Genetic analysis of winter hardiness in domesticated x wild chickpea crosses. [c] Genetic analysis of vernalization response in domesticated x wild chickpea crosses. [d] Digital expression analysis of a core selection of breeding and germplasm lines of chickpea that differ in winter hardiness and vernalization. [e] Identification of the genes involved in the chickpea winter hardiness and vernalization and construction of gene network controlling these traits. [f] Assessing the phenotypic and genetic correlations between winter hardiness, vernalization response and Ascochyta blight response in chickpea. The complexity of the vernalization response and the inefficiency of our selection experiments (below) required quitting the work on ascochyta response in the framework of this project. Background to the subject: Since its introduction to the Palouse region of WA and Idaho, and the northern Great Plains, chickpea has been a spring rotation legume due to lack of winter hardiness. The short growing season of spring chickpea limits its grain yield and leaves relatively little stubble residue for combating soil erosion. In Israel, chilling temperatures limit pod setting in early springs and narrow the effective reproductive time window of the crop. Winter hardiness and vernalization response of chickpea alleles were lost due to a series of evolutionary bottlenecks; however, such alleles are prevalent in its wild progenitor’s genepool. Major conclusions, solutions, achievements: It appears that both vernalization response and winter hardiness are polygenic traits in the wild-domesticated chickpea genepool. The main conclusion from the fieldwork in Israel is that selection of domesticated winter hardy and vernalization responsive types should be conducted in late flowering and late maturity backgrounds to minimize interference by daylength and temperature response alleles (see our Plant Breeding paper on the subject). The main conclusion from the US winter-hardiness studies is that excellent lines have been identified for germplasm release and continued genetic study. Several of the lines have good seed size and growth habit that will be useful for introgressing winter-hardiness into current chickpea cultivars to develop releases for autumn sowing. We sequenced the transcriptomes and profiled the expression of genes in 87 samples. Differential expression analysis identified a total of 2,452 differentially expressed genes (DEGs) between vernalized plants and control plants, of which 287 were shared between two or more Cicer species studied. We cloned 498 genes controlling vernalization, named CVRN genes. Each of the CVRN genes contributes to flowering date advance (FDA) by 3.85% - 10.71%, but 413 (83%) other genes had negative effects on FDA, while only 83 (17%) had positive effects on FDA, when the plant is exposed to cold temperature. The cloned CVRN genes provide new toolkits and knowledge to develop chickpea cultivars that are suitable for autumn-sowing. Scientific & agricultural implications: Unlike the winter cereals (barley, wheat) or pea, in which a single allelic change may induce a switch from winter to spring habit, we were unable to find any evidence for such major gene action in chickpea. In agricultural terms this means that an alternative strategy must be employed in order to isolate late flowering – ascochyta resistant (winter types) domesticated forms to enable autumn sowing of chickpea in the US Great Plains. An environment was identified in U.S. (eastern Washington) where autumn-sown chickpea production is possible using the levels of winter-hardiness discovered once backcrossed into advanced cultivated material with acceptable agronomic traits. The cloned CVRN genes and identified gene networks significantly advance our understanding of molecular mechanisms underlying plant vernalization in general, and chickpea in particular, and provide a new toolkit for switching chickpea from a spring-sowing to autumn-sowing crop.

The study of taxonomic composition and ecological characteristics of wood species on devastated lands as a theoretical basis for the phytomelioration of environment remains relevant nowadays. It was discovered 32 species, 25 genera and 15 families in the course of the analysis of woody plant community from devastated lands of Petrovsky waste rock dumps. Among them, allochthonous species (59.38%) have an advantage over autochthonous (40.63%) according to the quantitative indicators. It was established, hemiapophytes predominate among apophytesspecies andneophytes predominate among anthropophytesaccording to the time of entry, according to the method of invasion ergasiophytes, according to the degree of adaptation ergasiophytes and agriophytes. The results of our analysis indicate that the investigated woody plant communitycorresponds to the conditions of localization.

specific objectives of this proposal were to: 1) determine the location, topology, and oligomerization of FtsH11 protease; 2) identify the substrate/s of FtsH11 and the downstream components involved in maintaining thermostability of chloroplasts; 3) identify new elements involved in FtsH11 protease regulatory network related to HT adaptation processes in chloroplast; 4) Study the role of FtsH11 homologs from crop species in HT tolerance. Background to the topic: HT-tolerant varieties that maintain high photosynthetic efficiency at HT, and cope better with daily and seasonal temperature fluctuations are in great need to alleviate the effect of global warming on food production. Photosynthesis is a very complex process requiring accurate coordination of many complex systems and constant adjustments to the changing environments. Proteolytic activities mediated by various proteases in chloroplast are essential part of this process and critical for maintaining normal chloroplast functions under HT. However, little is known about mechanisms that contribute to adaptation of photosynthetic processes to HT. Our study has shown that a chloroplast-targeted Arabidopsis FtsH11 protease plays an essential and specific role in maintaining thermostability of thylakoids and normal photosynthesis at moderate HT. We hypothesized that FtsH11 homologs recently identified in other plant species might have roles similarly to that of AtFtsH1. Thus, dissecting the underlying mechanisms of FtsH11 in the adaptation mechanisms in chloroplasts to HT stress and other elements involved will aid our effort to produce more agricultural products in less favorable environments. Major conclusions, solutions, achievements - Identified the chloroplast inner envelope membrane localization of FtsH11. - Revealed a specific association of FtsH11 with the a and b subunits of CPN60. - Identified the involvement of ARC6, a protein coordinates chloroplast division machineries in plants, in FtsH11 mediated HT adaptation process in chloroplast. -Reveal possible association of a polyribonucleotide nucleotidyltransferase (cpPNPase), coded by At3G03710, with FtsH11 mediated HT adaptation process in chloroplast. - Mapped 4 additional loci in FtsH11 mediated HT adaptation network in chloroplast. - Demonstrated importance of the proteolytic activity of FtsH11 for thermotolerance, in addition to the ATPase activity. - Demonstrated a conserved role of plant FtsH11 proteases in chloroplast biogenesis and in maintaining structural and functional thermostability of chloroplast at elevated temperatures. Implications, both scientific and agricultural:Three different components interacting with FtsH11 were identified during the course of this study. At present, it is not known whether these proteins are directly involved in FtsH11mediated thermotolerance network in chloroplast and/or how these elements are interrelated. Studies aiming to connect the dot among biological functions of these networks are underway in both labs. Nevertheless, in bacteria where it was first studied, FtsH functions in heat shock response by regulating transcription level of σ32, a heat chock factor regulates HSPsexpression. FtsH also involves in control of biosynthesis of membrane components and quality control of membrane proteins etc. In plants, both Arc 6 and CPN60 identified in this study are essential in chloroplast division and developments as mutation of either one impairs chloroplast division in Arabidopsis. The facts that we have found the specific association of both α and β CPN60 with FtsH11 protein biochemically, the suppression/ enhancement of ftsh11 thermosensitive phenotype by arc6 /pnp allele genetically, implicate inter-connection of these networks via FtsH11 mediated network(s) in regulating the dynamic adaptation processes of chloroplast to temperature increases at transcriptional, translational and post-translational levels. The conserved role of FtsH11 proteases in maintaining thermostability of chloroplast at HT demonstrated here provides a foundation for improving crop photosynthetic performance at high temperatures.

Major threats to agricultural sustainability in the 21st century are drought, increasing temperatures, soil salinity and soilborne pathogens, all of which are being exacerbated by climate change and pesticide abolition and are burning issues related to agriculture in the Middle East. We have found that Class 2 fungal endophytes adapt native plants to environmental stresses (drought, heat and salt) in a habitat-specific manner, and that these endophytes can confer stress tolerance to genetically distant monocot and eudicot hosts. In the past, we generated a uv non-pathogenic endophytic mutant of Colletotrichum magna (path-1) that colonized cucurbits, induced drought tolerance and enhanced growth, and protected 85% - 100% against disease caused by certain pathogenic fungi. We propose: 1) utilizing path-1 and additional endophtyic microorganisms to be isolated from stress-tolerant local, wild cucurbit watermelon, Citrulluscolocynthis, growing in the Dead Sea and Arava desert areas, 2) generate abiotic and biotic tolerant melon crop plants, colonized by the isolated endophytes, to increase crop yields under extreme environmental conditions such as salinity, heat and drought stress, 3) manage soilborne fungal pathogens affecting curubit crop species growing in the desert areas. This is a unique and novel "systems" approach that has the potential to utilize natural plant adaptation for agricultural development. We envisage that endophyte-colonized melons will eventually be used to overcome damages caused by soilborne diseases and also for cultivation of this crop, under stress conditions, utilizing treated waste water, thus dealing with the limited resource of fresh water.