Elymus nutans genes for seed shattering and candidate gene-derived EST-SSR markers for germplasm evaluation.

BACKGROUND: Elymus nutans and E. sibiricus are two important forage grasses of the genus Elymus. But they are difficult to grow for commercial seed production due to serious seed shattering. We conducted a comparative transcriptome analysis of abscission zone to find possible transcription changes associated with seed shattering, explore candidate genes involved in seed shattering and identify candidate gene-based EST-SSR markers for germplasm evaluation. RESULTS: cDNA libraries from abscission zone (AZ) and non-abscission zone (NAZ) tissues of E. nutans were constructed and sequenced. A total of 111,667 unigenes were annotated and 7644 differentially expressed transcripts (DETs) were predicted, corresponding to 6936 up-regulated in AZ and 708 down-regulated in NAZ. We identified 489 candidate genes related to transcription factor, cell wall hydrolysis or modification, hydrolase activity, phytohormone signaling and response, lignin biosynthesis, and signal transduction or protein turnover. Eleven similar candidate genes involved in polygalacturonase activity, hydrolase activity, and mitogen-activated protein kinase were up-regulated in the abscission zone of the two Elymus species, suggesting these genes may have specific function for abscission zone development and seed shattering. A total of 67 polymorphic EST-SSR markers were developed and characterized based on the sequences of these candidate genes. Fourteen polymorphic EST-SSR primers were finally used to study genetic diversity in 48 E. nutans genotypes with contrasting seed shattering habit. The dendrogram based on molecular data showed that most accessions with similar seed shattering degree tended to group together. CONCLUSIONS: The expression data generated from this study provides an important resource for future molecular biological research. Many DETs were associated with abscission zone development, and EST-SSR loci related to candidate genes may have potential application in identifying trait-associated markers in E. nutans in the future.

Transcriptome profiling of Elymus sibiricus, an important forage grass in Qinghai-Tibet plateau, reveals novel insights into candidate genes that potentially connected to seed shattering.

BACKGROUND: Elymus sibiricus is an important forage grass in semi-arid regions, but it is difficult to grow for commercial seed production due to high seed shattering. To better understand the underlying mechanism and explore the putative genes related to seed shattering, we conducted a combination of morphological, histological, physiochemical and transcriptome analysis on two E. sibiricus genotypes (XH09 and ZhN03) that have contrasting seed shattering. RESULTS: The results show that seed shattering is generally caused by a degradation of the abscission layer. Early degradation of abscission layers was associated with the increased seed shattering in high seed shattering genotype XH09. Two cell wall degrading enzymes, cellulase (CE) and polygalacturonase (PG), had different activity in the abscission zone, indicating their roles in differentiation of abscission layer. cDNA libraries from abscission zone tissue of XH09 and ZhN03 at 7 days, 21 days and 28 days after heading were constructed and sequenced. A total of 86,634 unigenes were annotated and 7110 differentially expressed transcripts (DETs) were predicted from "XH09-7 vs ZhN03-7", "XH09-21 vs ZhN03-21" and "XH09-28 vs ZhN03-28", corresponding to 2058 up-regulated and 5052 down-regulated unigenes. The expression profiles of 10 candidate transcripts involved in cell wall-degrading enzymes, lignin biosynthesis and phytohormone activity were validated using quantitative real-time PCR (qRT-PCR), 8 of which were up-regulated in low seed shattering genotype ZhN03, suggesting these genes may be associated with reduction of seed shattering. CONCLUSIONS: The expression data generated in this study provides an important resource for future molecular biological research in E. sibiricus.

Elymus repens biomass allocation and acquisition as affected by light and nutrient supply and companion crop competition.

BACKGROUND AND AIMS: Competitive crops are a central component of resource-efficient weed control, especially for problematic perennial weeds such as Elymus repens Competition not only reduces total weed biomass, but denial of resources can also change the allocation pattern - potentially away from the underground storage organs that make perennial weeds difficult to control. Thus, the competition mode of crops may be an important component in the design of resource-efficient cropping systems. Our aim was to determine how competition from companion crops with different modes of competition affect E. repens biomass acquisition and allocation and discuss that in relation to how E. repens responds to different levels of light and nutrient supply. METHODS: Greenhouse experiments were conducted with E. repens growing in interspecific competition with increasing density of perennial ryegrass or red clover, or growing at three levels of both light and nutrient supply. KEY RESULTS: Elymus repens total biomass decreased with increasing biomass of the companion crop and the rate of decrease was higher with red clover than with perennial ryegrass, particularly for E. repens rhizome biomass. A reduced nutrient supply shifted E. repens allocation towards below-ground biomass while a reduced light supply shifted it towards shoot biomass. Red clover caused no change in E. repens allocation pattern, while ryegrass mostly shifted the allocation towards below-ground biomass, but the change was not correlated with ryegrass biomass. CONCLUSIONS: The companion crop mode of competition influences both the suppression rate of E. repens biomass acquisition and the likelihood of shifts in E. repens biomass allocation.

Variation in the Prevalence and Transmission of Heritable Symbionts Across Host Populations in Heterogeneous Environments.

Heritable microbes are abundant in nature and influential to their hosts and the communities in which they reside. However, drivers of variability in the prevalence of heritable symbionts and their rates of transmission are poorly resolved, particularly across host populations experiencing variable biotic and abiotic environments. To fill these gaps, we surveyed 25 populations of two native grasses (Elymus virginicus and Elymus canadensis) across the southern Great Plains (USA). Both grass species host heritable endophytic fungi (genus Epichloё) and can hybridize where their ranges overlap. From a subset of hosts, we characterized endophyte genotype using genetic loci that link to bioactive alkaloid production. First, we found mean vertical transmission rates and population-level prevalence were positively correlated, specifically for E. virginicus. However, both endophyte prevalence and transmission varied substantially across populations and did not strongly correlate with abiotic variables, with one exception: endophyte prevalence decreased as drought stress decreased for E. virginicus hosts. Second, we evaluated the potential influence of biotic factors and found that, after accounting for climate, endophyte genotype explained significant variation in symbiont inheritance. We also contrasted populations where host species co-occurred in sympatry vs. allopatry. Sympatry could potentially increase interspecific hybridization, but this variable did not associate with patterns of symbiont prevalence or transmission success. Our results reveal substantial variability in symbiont prevalence and transmission across host populations and identify symbiont genotype, and to a lesser extent, the abiotic environment as sources of this variation.

Change in abundance of three phytophagous mite species (Acari: Eriophyidae, Tetranychidae) on quackgrass in the presence of choke disease.

Phytophagous mites and endophytic fungi may interact when sharing a host plant, potentially influencing one another's growth or population dynamics; however, interactions between them are poorly known and remain largely unexplored. In this study, quantitative associations between three species of phytophagous mites and the endophytic fungus Epichloë bromicola Leuchtm. & Schardl (Clavicipitaceae, Ascomycotina) on quackgrass, Elymus repens (L.) Gould are reported. The mites' abundance was assessed on field-collected grass shoots that were either exhibiting choke disease symptoms or without the fungus. Overall, the abundance of Tetranychus urticae and Aculodes mckenziei was significantly lower on quackgrass plants infected by E. bromicola compared to plants without the fungus. Conversely, populations of Abacarus hystrix were significantly larger on plants colonised by the fungus than on uninfected plants. Thus, the presence of this endophytic fungus may have divergent effects on different phytophagous mite species although the basis of these effects is not yet known.

Breaking dormancy in freshly matured seeds of Elymus sibiricus, an important forage grass in the Tibetan Plateau.

Elymus sibiricus L. is an important forage grass widely distributed in Asia and is usually a dominant species on Tibetan Plateau alpine grasslands. Here, we used the seed development indices of 1000 seed weight, seed moisture content, and seed viability to compare the seed characteristics at 10, 20, 30, 40, 50, and 60 days after anthesis (DAA) in five populations of E. sibiricus growing in Gannan, China. Additionally, seeds collected at 60 DAA were air-dried for one month, and the primary germination percentage (GP) was determined in the laboratory. Treatment of seeds with 0.2% KNO3, 100 mg/L cytokinin, and 500 mg/L GA3 were tested for their effects on dormancy. A primary GP of 16% was found after 12 d of 15/25°C incubation, with no differences among the five populations. The 1000 seed weight and seed viability steadily increased and moisture content continuously fell with DAA. The optimal harvest time for E. sibiricus in an alpine pasture was 50 DAA. No effect on dormancy was found after treating seeds with 0.2% KNO3 or 100 mg/L cytokinin; however, a low concentration of GA3 induced a prompt and synchronized germination. Freshly matured E. sibiricus seeds were classified to be in non-deep physiologically dormant. Treatment of seeds with GA3 before sowing enhanced the emergence speed and seedling uniformity in E. sibiricus.

Indirect effects of an invasive annual grass on seed fates of two native perennial grass species.

Invasive plants exhibit both direct and indirect negative effects on recruitment of natives following invasion. We examined indirect effects of the invader Bromus tectorum (cheatgrass) on seed fates of two native grass species, Elymus elymoides and Pseudoroegneria spicata, by removing B. tectorum and by adding inoculum of the shared seed pathogen Pyrenophora semeniperda in factorial experiments at xeric and mesic field sites. We also included a supplemental watering treatment to increase emergence and also the potential for pathogen escape. We recorded emergence and survival of native seedlings and also determined the fate of unemerged seeds. At the xeric site, Pyrenophora-caused mortality was high (34%), and effects of other pathogens and failed emergence of germinants were smaller. Cheatgrass removal negatively affected both emergence (35 vs. 25%) and spring survival (69 vs. 42%). Pyrenophora-caused seed mortality increased with inoculum augmentation for both species (22 vs. 47% overall), but emergence was negatively impacted only for P. spicata (20 vs. 34%). At the mesic site, Pyrenophora-caused mortality was low (6%). Cheatgrass removal doubled emergence (26 vs. 14%). Seed mortality increased significantly with inoculum augmentation for P. spicata (12 vs. 5%) but not E. elymoides, while emergence was not significantly affected in either species. A large fraction of seeds produced germinants that failed to emerge (37%), while another large fraction (35%) was killed by other pathogens. We conclude that facilitation by cheatgrass at the xeric site but interference at the mesic site was probably mediated through litter effects that could be ameliorative or suppressive. Apparent competition between cheatgrass and native grasses could occur through Pyrenophora, especially in a xeric environment, but effects were weak or absent at emergence. This was probably because Pyrenophora attacks the same slow-germinating fraction that is subject to pre-emergence mortality from other causes, including attack by other pathogens such as Fusarium.

Fungal symbionts as manipulators of plant reproductive biology.

Symbioses have shaped the evolution of life, most notably through the fixation of heritable symbionts into organelles. The inheritance of symbionts promotes mutualism and fixation by coupling partner fitness. However, conflicts arise if symbionts are transmitted through only one sex and can shift host resources toward the sex through which they propagate. Such reproductive manipulators have been documented in animals with separate sexes but not in other phyla or sexual systems. Here we investigated whether the investment in male relative to female reproduction differed between hermaphroditic host plants with versus without a maternally inherited fungal symbiont. Plants with the fungus produced more seeds and less pollen than plants lacking the fungus, resulting in an ~40% shift in functional gender and a switch from male-biased to female-biased sex allocation. Given the ubiquity of endophytes in plants, reproductive manipulators of hermaphrodites may be widespread in nature.

Effects of plant age on antioxidant activity and endogenous hormones in Alpine Elymus sibiricus of the Tibetan Plateau.

Elymus sibiricus L. is a perennial forage species that has potential to serve as a forage source in livestock grazing systems. However, E. sibiricus has been shown to have a rapid and substantial reduction of aboveground biomass and seed yield after 3 or 4 years and an accelerated aging process. To determine possible aging mechanisms, we planted E. sibiricus seeds in triplicate blocks in 2012, 2015, and 2016, respectively, and harvested samples of leaves and roots at the jointing and heading stages in 2018 and 2019 to determine oxidative indices and endogenous hormones. The fresh aboveground biomass of 4- and 5-year old plants declined by 34.2% and 52.4% respectively compared with 3-year old plants, and the seed yield declined by 12.7% and 34.1%, respectively. The water content in leaves was 51.7%, 43.3%, and 35.6%, and net photosynthesis was 7.73, 6.35, and 2.08 µmol/m2·s in 3-, 4-, and 5-year old plants, respectively. The superoxide anion radical generation rate in leaves and roots did not show any aging pattern. There was a non-significant increase in malondialdehyde concentration with plant age, particularly in leaves and roots at the heading stage in 2019. The superoxide dismutase activity showed a declining trend with age of plant roots at the jointing stage in both 2018 and 2019. The peroxidase activity declined with plant age in both leaves and roots, for example, and the catalase activity in roots 4- and 7-year old plants declined by 13.8% and 0.85%, respectively, compared to 3-year old plants at the heading stage in 2018. Therefore, the reduced capacity of the antioxidant system may lead to oxidative stress during plant aging process. Overall, the concentrations of plant hormones, auxin (IAA), gibberellin (GA), zeatin (ZT), and abscisic acid (ABA) were significantly lower in roots than in leaves. The IAA concentration in leaves and roots exhibited different patterns with plant age. The ZT concentrations in leaves of 3-year old plants was 2.39- and 2.62-fold of those in 4- and 7-year old plants, respectively at the jointing stage, and in roots, the concentration declined with plant age. The changes in the GA concentration with plant age varied between the physiological stages and between years. The ABA concentrations appeared to increase with plant age, particularly in leaves. In conclusion, the aging process of E. sibiricus was apparently associated with an increase in oxidative stress, a decrease of ZT and an increase of ABA, particularly in roots. These findings highlight the effects of plant age on the antioxidant and endogenous hormone activity of E. sibiricus. However, these plant age-related trends showed variations between plant physiological stages and between harvest years that needs to be researched in the future to develop strategies to manage this forage species.

Facilitation and interference of seedling establishment by a native legume before and after wildfire.

In semi-arid ecosystems, heterogeneous resources can lead to variable seedling recruitment. Existing vegetation can influence seedling establishment by modifying the resource and physical environment. We asked how a native legume, Lupinus argenteus, modifies microenvironments in unburned and burned sagebrush steppe, and if L. argenteus presence facilitates seedling establishment of native species and the non-native annual grass, Bromus tectorum. Field treatments examined mechanisms by which L. argenteus likely influences establishment: (1) live L. argenteus; (2) dead L. argenteus; (3) no L. argenteus; (4) no L. argenteus with L. argenteus litter; (5) no L. argenteus with inert litter; and (6) mock L. argenteus. Response variables included soil nitrogen, moisture, temperature, solar radiation, and seedling establishment of the natives Elymus multisetus and Eriogonum umbellatum, and non-native B. tectorum. In both unburned and burned communities, there was higher spring soil moisture, increased shade and reduced maximum temperatures under L. argenteus canopies. Adult L. argenteus resulted in greater amounts of soil nitrogen (N) only in burned sagebrush steppe, but L. argenteus litter increased soil N under both unburned and burned conditions. Although L. argenteus negatively affected emergence and survival of B. tectorum overall, its presence increased B. tectorum biomass and reproduction in unburned plots. However, L. argenteus had positive facilitative effects on size and survival of E. multisetus in both unburned and burned plots. Our study indicates that L. argenteus can facilitate seedling establishment in semi-arid systems, but net effects depend on the species examined, traits measured, and level of abiotic stress.

Autotoxicity effect of water extracts from rhizosphere soil of Elymus sibiricus in different planting years on seed germination, physiological characteristics and phytohormones of seedlings.

Elymus sibiricus is a highly valuable perennial forage that is widely planted in the Qinghai-Tibet Plateau (QTP) region. However, E. sibiricus artificial grasslands have a short utilization lifespan, and reach the highest yield in the 2nd and 3rd year of plantation, then rapidly drop its productivity. We hypothesized that autotoxicity is one of the mechanisms for the reduction of the productivity. To test this hypothesis, we prepared the water extract from rhizosphere soils of E. sibiricus planted for 3, 4, 5, and 8 years and examined the effects of the extract concentrations at 0.05, 0.1, 0.2, and 0.5 g/mL on seed germination, seedling growth, physiological characteristics and phytohormones in the aboveground and roots of E. sibiricus. The results showed that the soil extract concentration, planting years, and their interaction had significant influences on the most of these indices. The soil extract inhibited the seed germination and growth of seedlings, and the inhibitory effects appeared to be stronger at the 0.5 g/mL rhizosphere soil extract for 5 and 8 years. The superoxide dismultase and peroxide activities, the free proline concentration, soluble sugar concentration were altered. The malondialdehyde concentration was, in general, increased, especially in 8 years soil extract. The indole acetic acid and gibberellic acids concentrations were lowered, while the abscisic acid concentration varied. These changes were depending on the extract concentration and the years of planting, without clear patterns in some of them in response to the extract concentration and planting years. In summary, autotoxicity can be a contributor to the retardation of the growth and development of artificial E. sibiricus grasslands. The inhibitory effects could be attribute to impaired antioxidant capacity and disturbance of osmortic-regulatory substances and plant hormones, and are more profound on the root than on the aboveground part of the seedlings.

Transcriptome profiles identify the common responsive genes to drought stress in two Elymus species.

Elymus, the largest genus of the Triticeae Dumort, is a forage grass in the Qinghai-Tibetan Plateau, where the climate has gradually become increasingly dry in recent years. To understand the mechanisms of the response to drought stress in Elymus species, we first investigated physiological and biochemical responses to polyethylene glycol (PEG-6000) simulated drought stress in two Elymus species, Elymus nutans and Elymus sibiricus, and found that E. nutans was more tolerant to drought stress than E. sibiricus. De novo transcriptome analysis of these two Elymus species treated with or without 10 % PEG-6000 revealed that a total of 1695 unigenes were commonly regulated by drought treatment in these two Elymus species, with 1614 unigenes up-regulated and 81 unigenes down-regulated. The coexpressed differentially expressed genes (DEGs) were enriched in regulation of transcription and gene expression in the GO database. KEGG pathway analysis indicated plant hormone signaling transduction were mostly enriched in co-expressed DEGs. Furthermore, genes annotated in the plant hormone signaling transduction were screened from co-expressed DEGs, and found that abscisic acid plays the major role in the drought stress tolerance of Elymus. Meanwhile, transcription factors screened from co-expressed DEGs were mainly classified into the ERF subfamily and WRKY, DREB, and HSF family members. Our results provide further reference for studying the response mechanism and culturing highly tolerant grasses of the Elymus species under drought stress.

Senescence in growth and reproductive allocation in a bunchgrass.

Senescence is a puzzling phenomenon. Few convincing studies of senescence in perennial herbaceous plants exist. While ramets are known to senesce, whether senescence of bunchgrasses actually occurs is not clear. In this study, we grew a set of plants of Elymus excelsus, a bunchgrass, to examine plant size, sexual reproduction and bud formation in individual plants in relation to their gradual ageing, in order to determine whether E. excelsus experiences senescence. We collected data in two consecutive years (2009 and 2010) from field samples of plants from 1 to 5 years old. Using regression models, we performed age-related analyses of growth and reproduction parameters. Our results showed that individual plant size (diameter, individual biomass), total biomass of ramets, number and biomass of reproductive ramets, percentage of ramets that were reproductive, reproductive allocation, over-wintering buds and juvenile ramets all declined with age. However, vegetative growth (number and biomass of vegetative ramets) did not decrease with age. Those plants that survived, dwindled in size as they aged. However, no plants shifted their resource allocation between growth and reproduction as they aged, so the shift in allocation did not account for the fall in size.

Consequences of asynchronous heading in a perennial bunchgrass (Elymus excelsus).

Reproduction is vital to plant population adaptation. The consequences of asynchronous reproduction in a perennial bunchgrass grass is not well studied. The heading reproductive tillers from early to late forms a continuum due to asynchronous heading and flowering in Elymus excelsus population. In two peak years of production, the three-year-old and four-year-old reproductive tillers of experimental E. excelsus population were marked from the early to late heading stage every four days at five different heading times and these tillers were harvested at the dough stage, respectively. The growth, biomass, seed production and reproductive allocation were measured to analyze the consequences of asynchronous reproduction. Reproductive tiller height, biomass, inflorescence length, inflorescence biomass, floret number, seed number, seed biomass, seed-set percentage, biomass allocation to inflorescence (RA1) and to seed (RA2) significantly decreased with the delay of heading date over the two years. Above ten phenotypic characteristics exponentially increased at a significant or extremely significant level with increasing differences in reproductive period. Reproductive tillers preferentially allocated the biomass to inflorescences, and then the inflorescences preferentially allocated the biomass to seeds throughout reproductive period. Earlier heading tillers had more contribution to E. excelsus population adaptation and development in the two peak years of production.

Novel fine-scale aerial mapping approach quantifies grassland weed cover dynamics and response to management.

Invasive weeds threaten the biodiversity and forage productivity of grasslands worldwide. However, management of these weeds is constrained by the practical difficulty of detecting small-scale infestations across large landscapes and by limits in understanding of landscape-scale invasion dynamics, including mechanisms that enable patches to expand, contract, or remain stable. While high-end hyperspectral remote sensing systems can effectively map vegetation cover, these systems are currently too costly and limited in availability for most land managers. We demonstrate application of a more accessible and cost-effective remote sensing approach, based on simple aerial imagery, for quantifying weed cover dynamics over time. In California annual grasslands, the target communities of interest include invasive weedy grasses (Aegilops triuncialis and Elymus caput-medusae) and desirable forage grass species (primarily Avena spp. and Bromus spp.). Detecting invasion of annual grasses into an annual-dominated community is particularly challenging, but we were able to consistently characterize these two communities based on their phenological differences in peak growth and senescence using maximum likelihood supervised classification of imagery acquired twice per year (in mid- and end-of season). This approach permitted us to map weed-dominated cover at a 1-m scale (correctly detecting 93% of weed patches across the landscape) and to evaluate weed cover change over time. We found that weed cover was more pervasive and persistent in management units that had no significant grazing for several years than in those that were grazed, whereas forage cover was more abundant and stable in the grazed units. This application demonstrates the power of this method for assessing fine-scale vegetation transitions across heterogeneous landscapes. It thus provides means for small-scale early detection of invasive species and for testing fundamental questions about landscape dynamics.

Plant reproduction is altered by simulated herbicide drift to constructed plant communities.

Herbicide drift may have unintended impacts on native vegetation, adversely affecting individual species and plant communities. To determine the potential ecological effects of herbicide drift, small plant community plots were constructed using 9 perennial species found in different Willamette Valley (OR, USA) grassland habitats. Studies were conducted at 2 Oregon State University (Corvallis, OR, USA) farms in 2 separate years, with single and combined treatments of 0.01 to 0.2× field application rates (f.a.r.) of 1119 g ha-1 for glyphosate (active ingredient [a.i.] of 830 g ha-1 acid glyphosate) and 560 g ha-1 a.i. for dicamba. Plant responses were percentage of cover, number of reproductive structures, mature and immature seed production, and vegetative biomass. Herbicide effects differed with species, year, and, to a lesser extent, farm. Generally, 0.1 to 0.2× f.a.r. of the herbicides were required to affect reproduction in Camassia leichtlinii, Elymus glaucus, Eriophyllum lanatum, Festuca idahoensis, Iris tenax, and Prunella vulgaris. Eriophyllum lanatum also had a significant increase in percentage of immature seed dry weight with 0.01× f.a.r. of dicamba or the combination of glyphosate plus dicamba. Other species showed similar trends, but fewer significant responses. These studies indicated potential effects of low levels of herbicides on reproduction of native plants, and demonstrated a protocol whereby species growing in a constructed plant community can be evaluated for ecological responses. Environ Toxicol Chem 2017;36:2799-2813. Published 2017 SETAC. This article is a US government work and, as such, is in the public domain in the United States of America.

Treatment of log yard run-off by irrigation of grass and willows.

Log yard run-off is a potential environmental risk, among other things because it creates an oxygen deficiency in receiving watercourses. This study was conducted to investigate the purification efficiency of soil-plant systems with couchgrass (Elymus repens) and willows (Salix sp.) when intensively irrigated with run-off from an open sprinkling system at a Norway spruce (Picea abies) log yard. The purification efficiency was determined both at the field scale (couchgrass) and in 68-L lysimeters (couchgrass and willows). Groundwater in the field and drainage water from the lysimeters were analysed for Total Organic Carbon (TOC), distillable phenols, total P, and total N. Retention of TOC, phenols and P occurred but no difference between couchgrass and willows was observed. The system had better purification capacity at the field scale than in the lysimeters.

Origin and Evolution of Allopolyploid Wheatgrass Elymus fibrosus (Schrenk) Tzvelev (Poaceae: Triticeae) Reveals the Effect of Its Origination on Genetic Diversity.

Origin and evolution of tetraploid Elymus fibrosus (Schrenk) Tzvelev were characterized using low-copy nuclear gene Rpb2 (the second largest subunit of RNA polymerase II), and chloroplast region trnL-trnF (spacer between the tRNA Leu (UAA) gene and the tRNA-Phe (GAA) gene). Ten accessions of E. fibrosus along with 19 Elymus species with StH genomic constitution and diploid species in the tribe Triticeae were analyzed. Chloroplast trnL-trnF sequence data suggested that Pseudoroegneria (St genome) was the maternal donor of E. fibrosus. Rpb2 data confirmed the presence of StH genomes in E. fibrosus, and suggested that St and H genomes in E. fibrosus each is more likely originated from single gene pool. Single origin of E. fibrosus might be one of the reasons causing genetic diversity in E. fibrosus lower than those in E. caninus and E. trachycaulus, which have similar ecological preferences and breeding systems with E. fibrosus, and each was originated from multiple sources. Convergent evolution of St and H copy Rpb2 sequences in some accessions of E. fibrosus might have occurred during the evolutionary history of this allotetraploid.

Diffusion limitations and metabolic factors associated with inhibition and recovery of photosynthesis following cold stress in Elymus nutans Griseb.

We studied the effects of cold stress (5°C) and re-warming (25°C) on gas exchange, photosystem II, key photosynthetic enzyme activities, gene expression, and carbohydrate metabolite concentrations in two Elymus nutans genotypes differing in cold resistance (DX, cold-tolerant and ZD, cold-sensitive). Cold stress led to irreversible reductions in photosynthetic rate. This reduction was accompanied by declining stomatal and mesophyll conductance (gs and gm), transpiration rate (Tr) and photochemical efficiency in both genotypes, however there were smaller decreases in DX than in ZD. Cold-tolerant DX maintained higher photosynthetic enzyme activities and transcript levels, as well as higher reducing sugar concentrations and sucrose accumulation. The relationship between Pn and internal leaf CO2 concentration (Pn/Ci curve) during cold and re-warming was analyzed to estimate the relative influence of stomatal and non-stomatal components on photosynthesis. Stomatal limitation, non-stomatal limitation, and CO2 compensation point (CP) increased in both genotypes under cold stress, but to a lesser extent in DX. Maximum CO2 assimilation rate (Pmax), and carboxylation efficiency (CE) declined, but DX had significantly higher levels of Pmax and CE than ZD. Following cold-stress recovery, the maximum quantum yield of PSII (Fv/Fm), apparent electron transport rate (ETR), Rubisco activity, Rubisco activation state and CE in DX resumed to the control levels. In contrast, Pn, Pmax, gs, gm, and Tr recovered only partially for DX, suggesting that incomplete recovery of photosynthesis in DX may be mainly related to diffusion limitations. Higher Rubisco large subunit (RbcL) and Rubisco activase (RCA) transcript levels, glyceraldehyde-3-phosphate dehydrogenase (GAPDH) activity, and carbohydrate accumulation contributed to higher photosynthetic recovery in DX. These results indicate that the maintenance of higher Pn and Pmax under cold stress and recovery in cold-tolerant DX could be attributed to reduced diffusion limitations and rapid recuperation of metabolic factors.

[Reproductive tillers phenotypic plasticity of Elymus excelsus population at different earring time in Songnen Plain of Northeast China].

The reproductive tillers of Elymus excelsus population in Songnen Plain were randomly marked with labels every other four days (five times in total) at early earring stage, and the marked tillers were harvested at dough stage, with the differences of their quantitative characteristics as well as the relationships between the quantitative characteristics and the shortened time of reproductive growth were analyzed. With the shortened time of reproductive growth, the quantitative characteristics of the reproductive tillers decreased gradually, which was not beneficial to the reproductive growth, reproductive allocation, and fruiting of reproductive tillers. The later the earring time, the larger the impact on fruiting and reproductive allocation was. When the reproductive growth time was shortened by 16 days, the grain biomass, setting penentage, reproductive allocation I , and reproductive allocation II decreased by 99.4%, 98.8%, 54.3%, and 99.2%, respectively. With the shortened time of reproductive growth, the tiller height decreased linearly, spike biomass, reproductive allocation I, and setting penentage decreased exponentially, tiller biomass, spike length, and floret number decreased powerly, and grain biomass, grain number, and reproductive allocation II decreased logarithmically.