Do invasive plants structure microbial communities to accelerate decomposition in intermountain grasslands?

Invasive plants are often associated with greater productivity and soil nutrient availabilities, but whether invasive plants with dissimilar traits change decomposer communities and decomposition rates in consistent ways is little known. We compared decomposition rates and the fungal and bacterial communities associated with the litter of three problematic invaders in intermountain grasslands; cheatgrass (Bromus tectorum), spotted knapweed (Centaurea stoebe) and leafy spurge (Euphorbia esula), as well as the native bluebunch wheatgrass (Pseudoroegneria spicata). Shoot and root litter from each plant was placed in cheatgrass, spotted knapweed, and leafy spurge invasions as well as remnant native communities in a fully reciprocal design for 6 months to see whether decomposer communities were species‐specific, and whether litter decomposed fastest when placed in a community composed of its own species (referred to hereafter as home‐field advantage–HFA). Overall, litter from the two invasive forbs, spotted knapweed and leafy spurge, decomposed faster than the native and invasive grasses, regardless of the plant community of incubation. Thus, we found no evidence of HFA. T‐RFLP profiles indicated that both fungal and bacterial communities differed between roots and shoots and among plant species, and that fungal communities also differed among plant community types. Synthesis. These results show that litter from three common invaders to intermountain grasslands decomposes at different rates and cultures microbial communities that are species‐specific, widespread, and persistent through the dramatic shifts in plant communities associated with invasions.     

Ecological genetics of vernalization response in Bromus tectorum L. (Poaceae)

BACKGROUND AND AIMS: Bromus tectorum (cheatgrass or downy brome) is an exotic annual grass that is dominant over large areas of former shrubland in western North America. To flower in time for seed production in early summer, B. tectorum plants generally require vernalization at winter temperatures, either as imbibed seeds or as established seedlings. METHODS: Variation in response to increasing periods of vernalization as seeds or seedlings for progeny of ten full-sib families from each of four B. tectorum populations from contrasting habitats was studied. KEY RESULTS: As vernalization was increased from 0 to 10 weeks, the proportion of plants flowering within 20 weeks increased, weeks to initiation of flowering decreased, and seed yield per plant increased, regardless of whether plants were vernalized as seeds or seedlings. Most of the variation was accounted for by differences among populations. Plants of the warm desert population flowered promptly even without vernalization, while those of the cold desert, foothill and montane populations showed incremental changes in response variables as a function of vernalization period. Populations differed in among-family variance, with the warm desert population generally showing the least variance and the cold desert population the most. Variation among populations and among families within populations decreased as vernalization period increased, whereas the non-genetic component of variance showed no such pattern. CONCLUSIONS: Variation in vernalization response was found to be adaptively significant and apparently represents the result of contrasting selection regimes on a range of founder genotypes.     

Estimating soil carbon sequestration under elevated CO2 by combining carbon isotope labelling with soil carbon cycle modelling

Elevated CO₂ concentrations generally stimulate grassland productivity, but herbaceous plants have only a limited capacity to sequester extra carbon (C) in biomass. However, increased primary productivity under elevated CO₂ could result in increased transfer of C into soils where it could be stored for prolonged periods and exercise a negative feedback on the rise in atmospheric CO₂. Measuring soil C sequestration directly is notoriously difficult for a number of methodological reasons. Here, we present a method that combines C isotope labelling with soil C cycle modelling to partition net soil sequestration into changes in new C fixed over the experimental duration (C_new) and pre‐experimental C (C_old). This partitioning is advantageous because the C_new accumulates whereas C_old is lost in the course of time (ΔC_new>0 whereas ΔC_old<0). We applied this method to calcareous grassland exposed to 600 μL CO₂ L^?1 for 6 years. The CO₂ used for atmospheric enrichment was depleted in ¹³C relative to the background atmosphere, and this distinct isotopic signature was used to quantify net soil C_new fluxes under elevated CO₂. Using ¹³C/¹²C mass balance and inverse modelling, the Rothamsted model ‘RothC’ predicted gross soil C_new inputs under elevated CO₂ and the decomposition of C_old. The modelled soil C pools and fluxes were in good agreement with experimental data. C isotope data indicated a net sequestration of ≈90 g C_new m^?2 yr^?1 in elevated CO₂. Accounting for C_old‐losses, this figure was reduced to ≈30 g C m^?2 yr^?1 at elevated CO₂; the elevated CO₂‐effect on net C sequestration was in the range of≈10 g C m^?2 yr^?1. A sensitivity and error analysis suggests that the modelled data are relatively robust. However, elevated CO₂‐specific mechanisms may necessitate a separate parameterization at ambient and elevated CO₂; these include increased soil moisture due to reduced leaf conductance, soil disaggregation as a consequence of increased soil moisture, and priming effects. These effects could accelerate decomposition of C_old in elevated CO₂ so that the CO₂ enrichment effect may be zero or even negative. Overall, our findings suggest that the C sequestration potential of this grassland under elevated CO₂ is rather limited.     

Invasive grass reduces aboveground carbon stocks in shrublands of the Western US

Understanding the terrestrial carbon budget, in particular the strength of the terrestrial carbon sink, is important in the context of global climate change. Considerable attention has been given to woody encroachment in the western US and the role it might play as a carbon sink; however, in many parts of the western US the reverse process is also occurring. The conversion of woody shrublands to annual grasslands involves the invasion of non-native cheatgrass ( Bromus tectorum ) which in turn leads to increased frequency and extent of fires. We compared carbon storage in adjacent plots of invasive grassland and native shrubland. We scaled-up the impact of this ecosystem shift using regional maps of the current invasion and of the risk of future invasion. The expansion of cheatgrass within the Great Basin has released an estimated 8±3 Tg C to the atmosphere, and will likely release another 50±20 Tg C in the coming decades. This ecosystem conversion has changed portions of the western US from a carbon sink to a source, making previous estimates of a western carbon sink almost certainly spurious. The growing importance of invasive species in driving land cover changes may substantially change future estimates of US terrestrial carbon storage.     

Interaction between the endophytic fungus Epichloe bromicola and the grass bromus erectus: effects of endophyte infection, fungal concentration and environment on grass growth and flowering

Epichloe bromicola is an endophytic fungal species that systemically and perennially colonizes intercellular spaces of leaf blades, leaf sheaths and culms of Bromus grass species. E. bromicola causes choke disease in B. erectus, suppressing maturation of most, if not all, host inflorescences. In an investigation of the interaction between fungus and host, we used a quantitative polymerase chain reaction technique to estimate the amount of fungal DNA, and thereby fungal concentration, in host plants. Fungal concentration was directly correlated with vegetative vigour of the plant, as measured by longest leaf length, number of tillers and vegetative above-ground biomass, suggesting that, during vegetative growth, the endophytic fungus is most beneficial for the plant when present in high concentrations. In contrast, the reproduction of the plant, as measured by the number of functional inflorescences, was inversely correlated with fungal concentration: the majority of infected plants, and all that were associated with high concentrations of fungi, were diseased. Thus, the benefit of endophyte infection for the plant is coupled with the disadvantages of infertility. Fungal concentration was shown to be at least in part genetically determined because fungal concentration differed significantly in different plant-endophyte genotype combinations (symbiotum). In a field experiment with normal and CO2-enriched environments, elevated CO2 levels favoured fungal reproductive vigour over host reproductive vigour, suggesting that these plant endophytes would be at a selective advantage in a corresponding environmental-change scenario. We conclude that a dynamic and complex relationship between fungal endophyte infection, fungal concentration, genotype and environment affects growth and fecundity of B. erectus and should contribute to the evolution of these plant-fungal interactions.     

Differential responses to nitrogen fertilization in native shrubs and exotic annuals common to mediterranean coastal sage scrub of California

This study examined the growth responses of exotic annuals and native shrubs to elevated N levels to test the hypothesis that increased N availability favors nitrophilous annuals over the slower-growing shrubs. The vegetation structure of the coastal sage scrub ecosystems in southern California is shifting from shrubland to annual grasslands. Over the last 30 years large tracts of wildlands, particularly those adjacent to urban centers, have lost significant native shrub cover, which has been replaced by exotic annuals native to the Mediterranean Basin. During this same time, air pollution has led to increased terrestrial eutrophication by atmospheric deposition. Changes in vegetation are often the result of changes in resource availability. The results of our experiments showed the three native shrubs tested to be more nitrophilous than the three annuals tested, which contrasts with most models of perennial species' adaptation to stressful environments. Under greenhouse conditions the annual grasses exhibited yield depression at the highest N treatments of 80 g g^–1 in soil. The three shrub species evaluated continued to increase shoot biomass at 80 g g^–1 N in soil. The grasses also exhibited increased tissue N concentrations with increased soil N in contrast with the shrubs where there was little difference in tissue N concentrations with increasing availability. Although the differential yield responses to elevated N do not explain the success of the annual vegetation in replacing shrubs, the inability of the shrubs to regulate growth under elevated N levels may explain the poor survival of mature individuals.     

盐胁迫对两种无芒雀麦种子萌发的影响

 在实验室中对两种无芒雀麦本地种（Bromus inermis cv. Xilinguole）和引进种 (Bromus stamineus)种子在浓度为0.5%、1.0%、1.5%的Na₂SO₄、NaCl和MgCl₂ 3种单盐胁迫下进行发芽实验，结果表明：与对照相比，在以上3种盐胁迫下，随着胁迫浓度的增加，两种无芒雀麦种子的发芽率、发芽指数和发芽值均有不同程度的降低，相对盐害率升高，种子开始发芽的时间推迟且其发芽过程延长；在3种盐中，NaCl的胁迫对两种植物种子萌发的抑制作用最大，其中浓度为1.0% NaCl的胁迫对引进种和本地种无芒雀麦发芽率的抑制分别达到了68.67%和14%，在浓度为1.5%的NaCl胁迫下两种供试植物的发芽率均为零；通过建立不同盐分处理与两种无芒雀麦种子发芽率之间的回归方程得出，引进无芒雀麦种子芽期的耐盐性较本地无芒雀麦强。     

Emergence of temperate pasture grases from different sowing depths: importance of seed weight, coleoptiele plus mesocotyl length and shoot strength

Inter-specific relationships between mean seed weight, coleoptile + mesocotyl (sub-coleoptile internode) length and width, shoot strength and emergence from different sowing depths were examined for timothy (Phleum pratense L., mean seed weight 0.33-0.48 mg depending on seed line), cocksfoot (Dactylis glomerata L., 0.65-0.78 mg), perennial ryegrass (Lolium perenne L., 1.71-2.19 mg), tall fescue (Festuca arundinucea Schreb., 1.73–2.60 mg), annual ryegrass (Lolium multiflorum Lam., 5.10-5.20 mg) and prairie grass (Bromus willdenowü Kunth., 10.5–12.2 mg). Across species at 10,30 and 60 mm sowing depths in the field and 10, 15 and 30 mm sowing depths under controlled environment conditions, there was a significant (P < 0.05) positive correlation between emergence % and mean seed weight. Across species at 10–30 mm sowing depth under controlled environment conditions, emergence % was not significantly correlated with coleoptile + mesocotyl length but there were significant positive correlations between emergence % and coleoptile and mesocotyl width: shoot strength increased with increased coleoptile width across species. For seed lines of timothy of different mean seed weight (0.21-0.81 mg), emergence %, coleoptile + mesocotyl length and coleoptile and mesocotyl width increased with increased seed weight at 10 and 15 mm sowing depth. Shoot strength increased with increased coleoptile width for timothy. For emerged and non-emerged cocksfoot and timothy seedlings regardless of seed weight, mean coleoptile + mesocotyl length was > 10 mm at 10 mm sowing depth. It is concluded that at 10–30 mm sowing depth, increased emergence % with increased seed weight across species is not due to increased coleoptile + mesocotyl length. It is proposed that increased emergence % with increased seed weight across species at 10–30 mm sowing depths and across seed lines of timothy at 10 mm sowing depth is primarily due to increased coleoptile and mesocotyl width resulting in increased shoot strength and hence an increased ability to penetrate the substrate.     

Ecological genetics of Bromus tectorum

Summary By incorporating demographic analyses of fitness components (e.g., survival and reproduction) within a reciprocal sowing design, we tested for 3 consecutive years whether local adaptation has occurred in the alien grass Bromus tectorum (cheatgrass) within 7 habitats along an environmental gradient from arid steppe to subalpine forest in the Intermontain Region of western North America. Patterns of emergence and survival were strongly influenced by the local environment. In terms of survival, expression of significant local adaptation in Tsuga heterophylla habitat varied among years. In contrast, relative differences in flowering time among seed sources were stable across sites and years. Populations from the arid steppe were the earliest to flower; flowering was latest in populations from the mesic Tsuga heterophylla habitat. In terms of net reproductive rate, evidence for local adaptation in B. tectorum was obtained in populations from habitats representing environmental extremes: an arid, saline site dominated by the shrub Sarcobatus vermiculatus and clearings within the cool, mesic Tsuga heterophylla forest habitat. Unlike the plants introduced from other sites, members of the resident population at the Sarcobatus site flowered and produced seeds before soil water became limiting. In contrast, net reproductive rates in other habitats were sometimes the lowest for populations in their home site. This lack of an advantage for local populations within more environmentally moderate sites suggests that limited dispersal may restrict the rate at which superior genotypes are introduced into a particular site.     

The effect of shading on photosynthesis,growth, and regrowth following defoliation for Bromus tectorum

Summary The effect of full sunlight, 60%, or 90% attenuated light on photosynthetic rate, growth, leaf morphology, dry weight allocation patterns, phenology, and tolerance to clipping was examined in the glasshouse for steppe populations of the introduced grass, Bromus tectorum. The net photosynthetic response to light for plants grown in shade was comparable to responses for plants grown in full sunlight. Plants grown in full sunlight produced more biomass, tillers and leaves, and allocated a larger proportion of their total production to roots than plants grown in shade. The accumulation of root and shoot biomass over the first two months of seedling growth was primarily responsible for the larger size at harvest of plants grown in full sunlight. Plants grown under 60% and 90% shade flowered an average of 2 and 6 weeks later, respectively, than plants grown in full sunlight. Regrowth after clipping was greater for plants grown in full sunlight compared to those grown in shade. Even a one-time clipping delayed flowering and seed maturation; the older the individual when leaf area was removed, the greater the delay in its phenology. Repeated removal of leaf area was more frequently fatal for plants in shade than in full sunlight. For plants originally grown in full sunlight, regrowth in the dark was greater than for shaded plants and was more closely correlated to non-flowering tiller number than to plant size. This correlation suggests that etiolated regrowth is more likely regulated by the number of functional meristems than by differences in the size of carbohydrate pools. Thus, shading reduces the rate of growth, number of tillers, and ability to replace leaf area lost to herbivory for B. tectorum. These responses, in turn, intensify the effect of competition and defoliation for this grass in forests. B. tectorum is largely restricted to forest gaps at least in part because of its inability to acclimate photosynthetically, the influence of shade on resource allocation, and the role of herbivory in exacerbating these effects.