Local‐scale biotic interactions embedded in macroscale climate drivers suggest Eltonian noise hypothesis distribution patterns for an invasive grass

A hierarchical view of niche relations reconciles the scale‐dependent effects of abiotic and biotic processes on species distribution patterns and underlies most current approaches to distribution modeling. A key prediction of this framework is that the effects of biotic interactions will be averaged out at macroscales – an idea termed the Eltonian noise hypothesis (ENH). We test this prediction by quantifying regional variation in local abiotic and biotic niche relations and assess the role of macroclimate in structuring biotic interactions, using a non‐native invasive grass, Microstegium vimineum, in its introduced range. Consistent with hierarchical niche relations and the ENH, macroclimate structures local biotic interactions, while local abiotic relations are regionally conserved. Biotic interactions suppress M. vimineum in drier climates but have little effect in wetter climates. A similar approach could be used to identify the macroclimatic conditions under which biotic interactions affect the accuracy of local predictions of species distributions.     

Latitudinal variation in plant–herbivore interactions in European salt marshes

Ecological interactions often vary geographically. Work in salt marshes on the Atlantic Coast of the United States has documented community-wide latitudinal gradients in plant palatability and plant traits that may be driven in part by greater herbivore pressure at low latitudes. To determine if similar patterns exist elsewhere, we studied six taxa of saltmarsh plants ( Atriplex , Juncus , Limonium , Salicornia , Spartina and Suaeda ) at European sites at high (Germany and the Netherlands) and low (Portugal and Spain) latitudes. We conducted feeding assays using both native and non-native consumers, and documented patterns of herbivore damage in the field. As in the United States, high-latitude plants tended to be more palatable than low-latitude plants when offered to consumers in paired feeding assays in the laboratory, although assays with grasshopper consumers were less consistent than those with crab consumers, and plants in the field at low-latitude sites tended to experience greater levels of herbivore pressure than plants at high-latitude sites. Similarly, high-latitude leaf litter was more palatable than litter from low-latitude plants when offered to consumers in paired feeding assays in the laboratory. Latitudinal gradients in plant palatability and herbivore pressure may be a general phenomenon, and may contribute to latitudinal gradients in decomposition processes.     

Manganese as an ecological factor in salt marshes

The plant available manganese concentration (Mn²⁺) of salt-marsh sediments was compared to that of acidic and neutral soils. The mean soil-manganese concentration was higher in the top 1 cm of salt-marsh soil than in the neutral soil and comparable to that of the acidic soil (0–5 cm). A peak in the soil-manganese concentration in the upper marsh was observed one week after the spring tide but this effect was not evident in the lower marsh. Despite these differences, there was no correlation between mean manganese concentration and position on the marsh.The response to manganese of salt-marsh halophytes was studied by measuring growth and root elongation in a range of Mn²⁺ concentrations with and without sodium chloride. Although there was a differential response to manganese between salt-marsh species, manganese resistance was not related to position on the marsh. Most of the species investigated were tolerant of Mn²⁺ at concentrations higher than normally recommended for plant growth. Moreover a salt-marsh ecotype of Festuca rubra was found to have a higher manganese resistance than an inland ecotype of the same species.When sodium chloride was included in the growth medium, salt-marsh plants had a greatly increased resistance to manganese associated with a reduced uptake. This effect is reflected in the tissue-manganese concentration which was lower than in Deschampsia flexuosa although both groups of plants were exposed to a similar range of Mn²⁺ concentrations. It is concluded that sodium chloride markedly reduces the phytotoxicity of manganese in salt marshes.Nomenclature following Clapham, Tutin & Warburg (1968). Flora of the British Isles.The work was carried out while one of us (C. E. Singer) was in receipt of an SERC studentship, which is gratefully acknowledged.     

Differential response to frequency-dependent interactions: an experimental test using genotypes of an invasive grass

Positive feedbacks have been suggested as a means for non-indigenous species to successfully invade novel environments. Frequency-dependent feedbacks refer to a species performance being dependent on its local abundance in the population; however, frequency dependence is often described as a monolithic trait of a species rather than examining the variation in response for individual genotypes and fitness traits. Here, we investigate frequency-dependent outcomes for individual genotypes and fitness-related traits for the invasive grass Phalaris arundinacea. We tested for competition-mediated frequency dependence by establishing hexagonal arrays with the center target plant surrounded by either same, different or no genotype neighbors to determine how changing the small-scale frequency neighborhood-influenced invasion success. We used a Bayesian ANOVA approach which allowed us to easily accommodate our non-normal dataset and found that same neighbor plots had greater biomass production than different neighbor plots. Target plants also had greater stem height and aboveground biomass when surrounded by same genotype neighbors. A greenhouse experiment did not support the hypothesis that increased mycorrhizal associations were the cause of positive frequency dependence. We devised a frequency-dependent metric to quantify the extent of fitness-related differences for individual genotypes and found that individual genotypes showed a range of both positive and negative responses to different frequency treatments; however, only positive responses were statistically significant. The small-scale genotypic neighborhood had no effect for the fitness-related traits of leaf number, belowground biomass and total biomass. We demonstrate that individual invasive genotypes respond differently to changing frequency neighborhoods and that growth responses do not respond with the same direction and magnitude. A range of frequency-dependent responses may allow genotypes to invade a wide range of environments.     

Functions of the respiratory burst oxidase in biotic interactions, abiotic stress and development

The production of reactive oxygen intermediates (ROI) is among the earliest temporal events following pathogen recognition in plants. Initially, ROI were thought to be cell-death executioners. Emerging evidence, however, suggests a broader role for ROI as signals that mediate responses to infection, the abiotic environment, developmental cues, and programmed cell death in different cell types. The Respiratory burst oxidase homolog (Rboh) gene family encodes the key enzymatic subunit of the plant NADPH oxidase. Rboh proteins are the source of ROI produced following pathogen recognition and in a variety of other processes.     

Adaptive phenotypic divergence in an annual grass differs across biotic contexts*

Climate is a powerful force shaping adaptation within species, yet adaptation to climate occurs against a biotic background: species interactions can filter fitness consequences of genetic variation by altering phenotypic expression of genotypes. We investigated this process using populations of teosinte, a wild annual grass related to maize (Zea mays ssp. mexicana), sampling plants from 10 sites along an elevational gradient as well as rhizosphere biota from three of those sites. We grew half‐sibling teosinte families in each biota to test whether trait divergence among teosinte populations reflects adaptation or drift, and whether rhizosphere biota affect expression of diverged traits. We further assayed the influence of rhizosphere biota on contemporary additive genetic variation. We found that adaptation across environment shaped divergence of some traits, particularly flowering time and root biomass. We also observed that different rhizosphere biota shifted expressed values of these traits within teosinte populations and families and altered within‐population genetic variance and covariance. In sum, our results imply that changes in trait expression and covariance elicited by rhizosphere communities could have played a historical role in teosinte adaptation to environments and that they are likely to play a role in the response to future selection.     

WsSGTL1 gene from Withania somnifera,modulates glycosylation profile,antioxidant system and confers biotic and salt stress tolerance in transgenic tobacco

Glycosylation of sterols, catalysed by sterol glycosyltransferases (SGTs), improves the sterol solubility, chemical stability and compartmentalization, and helps plants to adapt to environmental changes. The SGTs in medicinal plants are of particular interest for their role in the biosynthesis of pharmacologically active substances. WsSGTL1, a SGT isolated from Withania somnifera, was expressed and functionally characterized in transgenic tobacco plants. Transgenic WsSGTL1-Nt lines showed an adaptive mechanism through demonstrating late germination, stunted growth, yellowish-green leaves and enhanced antioxidant system. The reduced chlorophyll content and chlorophyll fluorescence with decreased photosynthetic parameters were observed in WsSGTL1-Nt plants. These changes could be due to the enhanced glycosylation by WsSGTL1, as no modulation in chlorophyll biogenesis-related genes was observed in transgenic lines as compared to wildtype (WT) plants. Enhanced accumulation of main sterols like, campesterol, stigmasterol and sitosterol in glycosylated form was observed in WsSGTL1-Nt plants. Apart from these, other secondary metabolites related to plant’s antioxidant system along with activities of antioxidant enzymes (SOD, CAT; two to fourfold) were enhanced in WsSGTL1-Nt as compared to WT. WsSGTL1-Nt plants showed significant resistance towards Spodoptera litura (biotic stress) with up to 27 % reduced larval weight as well as salt stress (abiotic stress) with improved survival capacity of leaf discs. The present study demonstrates that higher glycosylation of sterols and enhanced antioxidant system caused by expression of WsSGTL1 gene confers specific functions in plants to adapt under different environmental challenges.     

Latitudinal and climate-driven variation in the strength and nature of biological interactions in New England salt marshes

We examined the linkage between climate and interspecific plant interactions in New England salt marshes. Because harsh edaphic conditions in marshes can be ameliorated by neighboring plants, plant neighbors can have net competitive or facilitative interactions, depending on ambient physical stresses. In particular, high soil salinities, which are largely controlled by solar radiation and the evaporation of marsh porewater, can be ameliorated by plant neighbors under stressful conditions leading to facilitative interactions. Under less stressful edaphic conditions, these same neighbors may be competitors. In this paper, we use this mechanistic understanding of marsh plant interactions to examine the hypothesis that latitudinal and inter-annual variation in climate can influence the nature and strength of marsh plant species interactions. We quantified the relationship between climate and species interactions by transplanting marsh plants into ambient vegetation and unvegetated bare patches at sites north and south of Cape Cod, a major biogeographic barrier on the east coast of North America. We hypothesized that the cooler climate north of Cape Cod would lead to fewer positive interactions among marsh plants. We found both latitudinal and inter-annual variation in the neighbor relations of marsh plants that paralleled latitudinal differences in temperature and salinity. South of Cape Cod, plant neighbor interactions tended to be more facilitative, whereas north of Cape Cod, plant neighbor interactions were more competitive. At all sites, soil salinity increased and plant neighbor interactions were more facilitative in warmer versus cooler years. Our results show that interspecific interactions can be strikingly linked to climate, but also reveal that because the sensitivity of specific species interactions to climatic variation is highly variable, predicting how entire communities will respond to climate change will be difficult, even in relatively simple, well-studied systems.     

Pyrite accumulation in salt marshes in the Eastern Scheldt,southwest Netherlands

Pore water composition, pyrite distribution and pyrite crystal morphology of sediments from salt marshes in the Eastern Scheldt, southwestern Netherlands, were examined from July 1984 to October 1986.Hydrology and marsh vegetation were the chief determinants of pyrite accumulation. In the bare sediments of pans in the low marsh, highly reducing conditions prevailed just below the surface. At these sites, practically all the incoming detrital pyrite (0.5–1% FeS₂) was preserved. The in-situ formation of pyrites was negligible in these anoxic sediments.All incoming detrital pyrite was oxidized in the surface layers (0–10 cm) of the medium-high marsh overgrown withSpartina anglica. Pyrite was formed at a rate of 2.6–3.8 mol S-FeS₂m^–2yr^–1 in a narrow range of depths (15–20cm), at the interface of the oxidizing and underlying reducing sediment. At this interface the concentration profiles of Fe²⁺ and dissolved S intersected. The role of the rhizosphere is discussed in connection with pyrite formation. No further pyrite formation occurred deeper in the sediment. This resulted in the build up of high concentrations of dissolved S and acid volatile sulfides (AVS). The decrease with depth in oxalate-extractable Fe indicated that most of the iron oxyhydroxides (70–80%) had been transformed to pyrite. Another 10–20% of oxalate-extractable Fe was present as AVS. The abundance of framboidal pyrite particles and the high concentrations of AVS and dissolved S indicated that the formation of pyrite occurred via iron monosulfide intermediatesThere was a linear relationship between the organic carbon and the S-FeS₂ content in theSpartina overgrown reducing sediment. The mean C/S ratio was 4.2.     

Water-induced stress influences the relative investment in cleistogamous and chasmogamous flowers of an invasive grass,Microstegium vimineum (Poaceae)

Background: Global climate change has the potential to shape evolutionary trajectories of invasive species via many routes, including through changes in mating systems. Many cleistogamous (CL) plants adjust investment in CL (selfed) vs. chasmogamous (CH, potentially outcrossed) progeny across environmental gradients. However, the details of such adjustments are lacking for highly invasive plant species.

Aims: We used a highly invasive grass, Microstegium vimineum, as a model for understanding how changes in water-induced stress (including potential associated changes in soil nutrient availability) might affect mating systems and thus evolutionary change in invasive species. We predicted that plants would respond to increased water-induced stress through a relative reduction in investment in CL vs. CH reproduction (i.e., a decrease in the CL:CH ratio).

Methods: Under greenhouse conditions, we measured fecundity (number of inflorescences and florets per plant) as well as relative investment in CL vs. CH florets (CL:CH ratios for number of inflorescences, florets per inflorescence, overall florets) in response to three watering treatments approximating mesic (low) to inundated (high) conditions.

Results: Plant biomass was significantly lower in high-watering treatment relative to intermediate and low treatments, indicating that the high-water condition was stressful. Contrary to expectations, stressed plants significantly increased relative investment in CL reproduction, a pattern associated with decreased inflorescence number and increased numbers of CL florets per inflorescence.

Conclusions: We conclude that changes in water-induced stress could strongly influence realised rates of outcrossing in this invasive plant, leading to mating system evolution, and altered invasiveness.     

Evidence for rapid evolution of phenology in an invasive grass

Evolutionary dynamics of integrative traits such as phenology are predicted to be critically important to range expansion and invasion success, yet there are few empirical examples of such phenomena. In this study, we used multiple common gardens to examine the evolutionary significance of latitudinal variation in phenology of a widespread invasive species, the Asian short‐day flowering annual grass Microstegium vimineum. In environmentally controlled growth chambers, we grew plants from seeds collected from multiple latitudes across the species' invasive range. Flowering time and biomass were both strongly correlated with the latitude of population origin such that populations collected from more northern latitudes flowered significantly earlier and at lower biomass than populations from southern locations. We suggest that this pattern may be the result of rapid adaptive evolution of phenology over a period of less than one hundred years and that such changes have likely promoted the northward range expansion of this species. We note that possible barriers to gene flow, including bottlenecks and inbreeding, have apparently not forestalled evolutionary processes for this plant. Furthermore, we hypothesize that evolution of phenology may be a widespread and potentially essential process during range expansion for many invasive plant species.     

Alkali grass resists salt stress through high [K+] and an endodermis barrier to Na+

In order to understand the salt-tolerance mechanism of alkali grass (Puccinellia tenuiflora) compared with wheat (Triticum aestivum L.), [K(+)] and [Na(+)] in roots and shoots in response to salt treatments were examined with ion element analysis and X-ray microanalysis. Both the rapid K(+) and Na(+) influx in response to different NaCl and KCl treatments, and the accumulation of K(+) and Na(+) as the plants acclimated to long-term stress were studied in culture- solution experiments. A higher K(+) uptake under normal and saline conditions was evident in alkali grass compared with that in wheat, and electrophysiological analyses indicated that the different uptake probably resulted from the higher K(+)/Na(+) selectivity of the plasma membrane. When external [K(+)] was high, K(+) uptake and transport from roots to shoots were inhibited by exogenous Cs(+), while TEA (tetraethylammonium) only inhibited K(+) transport from the root to the shoot. K(+) uptake was not influenced by Cs(+) when plants were K(+) starved. It was shown by X-ray microanalysis that high [K(+)] and low [Na(+)] existed in the endodermal cells of alkali grass roots, suggesting this to be the tissue where Cs(+) inhibition occurs. These results suggest that the K(+)/Na(+) selectivity of potassium channels and the existence of an apoplastic barrier, the Casparian bands of the endodermis, lead to the lateral gradient of K(+) and Na(+) across root tissue, resulting not only in high levels of [K(+)] in the shoot but also a large [Na(+)] gradient between the root and the shoot.