Conservative ecological and evolutionary patterns in liverwort–fungal symbioses

Liverworts, the most ancient group of land plants, form a range of intimate associations with fungi that may be analogous to the mycorrhizas of vascular plants. Most thalloid liverworts contain arbuscular mycorrhizal glomeromycete fungi similar to most vascular plants. In contrast, a range of leafy liverwort genera and one simple thalloid liverwort family (the Aneuraceae) have switched to basidiomycete fungi. These liverwort switches away from glomeromycete fungi may be expected to parallel switches undergone by vascular plants that target diverse lineages of basidiomycete fungi to form ectomycorrhizas. To test this hypothesis, we used a cultivation-independent approach to examine the basidiomycete fungi associated with liverworts in varied worldwide locations by generating fungal DNA sequence data from over 200 field collections of over 30 species. Here we show that eight leafy liverwort genera predominantly and consistently associate with members of the Sebacina vermifera species complex and that Aneuraceae thalloid liverworts associate nearly exclusively with Tulasnella species. Furthermore, within sites where multiple liverwort species co-occur, they almost never share the same fungi. Our analyses reveal a strikingly conservative ecological and evolutionary pattern of liverwort symbioses with basidiomycete fungi that is unlike that of vascular plant mycorrhizas.     

Complexity in the eutrophication–harmful algal bloom relationship,with comment on the importance of grazing

This article seeks to guide the conceptual development and field application of the eutrophication–HAB hypothesis. After considering the evidence for this hypothesis, the importance of exogenous nutrients as a chemical habitat conditioner and the “family” of nutrient regulated effects that occur are discussed. The various definitions of eutrophication are applied; the conceptual ambiguity over how to perceive eutrophication, and the need to view eutrophication as a process and not as an ecological state are considered. The habitat irradiance-nutrient-flushing gradient regulates the bloom potential in response to exogenous nutrients. There is an apparent species-specific paradox within the eutrophication–HAB hypothesis related to the phycotoxin synthesis–nutrient relationship. Two nutrient–toxin relationships occur: toxin biosynthesis during nutrient sufficiency, and biosynthesis that requires a nutrient limitation. HAB events can be just as much nutrient depletion events as nutrient stimulated events, and whether the former develops depends upon the specific cellular toxicity–nutrient relationship of the bloom species. The importance of grazing in bloom regulation is highlighted. HABs and red tides generally should be viewed as blooms that are regulated by coupled nutrient-grazer processes – nutrient stimulation alone is inadequate, even when exogenous nutrients are not a factor. In assessing the eutrophication–HAB relationship, the collective grazing behavior of the micro-zooplankton, herbivorous copepods, filter feeding benthos, benthic larvae and, when present, omnivorous nekton must be considered. The importance of grazing in the bloom behavior of HAB species is illustrated using field and experimental data during a 5-month brown tide in Narragansett Bay. A cascade in grazing pressure regulated this bloom, with the initial collapse and then restoration of grazing pressure progressing through 7 stages of collective grazing pressure by micro-zooplankton, herbivorous copepods, benthic larvae, benthic filter feeders, and lytic virus infection.     

Pineal–adrenal–immune system relationship under thermal stress: effect on physiological,endocrine, and non-specific immune response in goats

The purpose of the investigation was to observe the pineal–adrenal–immune system relationships and their influence on non-specific immune response in female goats under short-term thermal stress. Six female goats had been exposed to 40°C and 60% relative humidity in the psychrometric chamber for 17 days. Blood samples were obtained on days 0 and 10 to establish control and thermal stress effects, respectively. Chemical adrenalectomy was achieved by injecting metyrapone (100 mg/kg body weight) followed by exogenous melatonin treatment (0.1 mg/kg body weight) from 11th to 17th day of experiment. Thermal stress significantly (P ≤ 0.05) altered the physiological responses. Metyrapone and melatonin treatment significantly (P ≤ 0.05) reduced the thermal-stress-induced increase in plasma concentrations of cortisol and corticosterone while significantly (P ≤ 0.05) increased the plasma melatonin on days 11 and 17. Furthermore, these treatments significantly (P < 0.05) increased the phagocytic activity of neutrophils as compared to both control and thermal exposure values from 11–17 days of experiment. The data generated from this study help us to understand the functional relationship between pineal, adrenal, and immune system, and how this relationship modifies the non-specific immune response for the well being of goats during thermal stress.     

Are patterns in the taxonomic,biological and ecological traits of water beetles congruent in Mediterranean ecosystems?

1. Coleoptera species show considerable diversity in life histories and ecological strategies, which makes possible their wide distribution in freshwater habitats, including highly stressed ones such as saline or temporary waterbodies. Explaining how particular combinations of traits allow species to occupy distinctive habitats is a central question in ecology. 2. A total of 212 sites, sampled over a wide range of inland aquatic habitats in the south‐eastern Iberian Peninsula, yielded 272 species belonging to 68 genera and 11 families. The affinities of genera for 11 biological and 11 ecological traits, gathered from literature and the authors’ own expertise, were used to assess the degree of congruence between taxonomic, biological and ecological traits. 3. Taxonomic richness was significantly related to the number of both biological and ecological trait categories, with the richest families also showing the highest functional and ecological diversity. A fuzzy correspondence analysis performed on the abundance‐weighed array of biological traits separated genera according to categories of diet, feeding habits, respiration, reproduction and locomotion. A similar analysis of ecological traits revealed that preferences related to longitudinal distribution (headwater to mouth), local habitat and current velocity best discriminated genera. At the family level, there was a distinctive functional grouping of genera based on biological traits. Only Elmidae showed noticeable homogeneity across genera for both biological and ecological traits. 4. Co‐inertia analysis demonstrated a significant match between biological and ecological traits (Rv‐correlation = 0.35, P < 0.001). Elmidae genera displayed the highest concordance, whereas Hydraenidae demonstrated the lowest. 5. These results indicate that the predominance of habitat filtering processes in headwater streams yields biological trait conservatism (as shown by Elmidae genera), as well as trait convergence for some specific traits (for instance, respiration) among certain Dytiscidae genera and other typical rheophilic taxa, whereas other biotic factors, such as competition among species, appear more prominent in less stressed habitats. Further knowledge of traits, especially regarding physiological capabilities, is needed to better understand water beetle life history strategies.     

Interpreting spatial patterns in redox and coupled water–nitrogen fluxes in the streambed of a gaining river reach

Water pathways through permeable riverbeds are multi-dimensional, including lateral hyporheic exchange flows as well as vertical (upwelling and downwelling) fluxes. The influence of different pathways of water on solute patterns and the supply of nitrate and other redox-sensitive chemical species in the riverbed is poorly understood but could be environmentally significant. For example, nitrate-rich upwelling water in the gaining reaches of groundwater-fed rivers has the potential to supply significant quantities of nitrate through the riverbed to surface waters, constraining opportunities to deliver the goals of the EU Water Framework Directive to achieve ‘good ecological status’. We show that patterns in porewater chemistry in the armoured river bed of a gaining reach (River Leith, Cumbria) reflect the spatial variability in different sources of water; oxic conditions being associated with preferential discharge from groundwater and reducing conditions with longitudinal and lateral fluxes of water due to water movement from riparian zones and/or hyporheic exchange flows. Our findings demonstrate the important control of both vertical and lateral water fluxes on patterns of redox-sensitive chemical species in the river bed. Furthermore, under stable, baseflow conditions (<Q₉₀) a zone of preferential discharge, comprising 20 % of the reach by area contributes 4–9 % of the total nitrate being transported through the reach in surface water, highlighting the need to understand the spatial distribution of such preferential discharge locations at the catchment scale to establish their importance for nitrate delivery to the stream channel.     

Leaf phosphorus influences the photosynthesis–nitrogen relation: a cross-biome analysis of 314 species

The ecophysiological linkage of leaf phosphorus (P) to photosynthetic capacity (A _max) and to the A _max–nitrogen relation remains poorly understood. To address this issue we compiled published and unpublished field data for mass-based A _max, nitrogen (N) and P (n = 517 observations) from 314 species at 42 sites in 14 countries. Data were from four biomes: arctic, cold temperate, subtropical (including Mediterranean), and tropical. We asked whether plants with low P levels have low A _max, a shallower slope of the A _max–N relationship, and whether these patterns have a geographic signature. On average, leaf P was substantially lower in the two warmer than in the two colder biomes, with the reverse true for N:P ratios. The evidence indicates that the response of A _max to leaf N is constrained by low leaf P. Using a full factorial model for all data, A _max was related to leaf N, but not to leaf P on its own, with a significant leaf N ×  leaf P interaction indicating that the response of A _max to N increased with increasing leaf P. This was also found in analyses using one value per species per site, or by comparing only angiosperms or only woody plants. Additionally, the slope of the A _max–N relationship was higher in the colder arctic and temperate than warmer tropical and subtropical biomes. Sorting data into low, medium, and high leaf P groupings also showed that the A _max–N slope increases with leaf P. These analyses support claims that in P-limited ecosystems the A _max–N relationship may be constrained by low P, and are consistent with laboratory studies that show P-deficient plants have limited ribulose-1,5-bisphosphate regeneration, a likely mechanism for the P influence upon the A _max–N relation. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Spatial patterns of photosynthesis in thin- and thick-leaved epiphytic orchids: unravelling C3–CAM plasticity in an organ-compartmented way

Background and Aims

A positive correlation between tissue thickness and crassulacean acid metabolism (CAM) expression has been frequently suggested. Therefore, this study addressed the question of whether water availability modulates photosynthetic plasticity in different organs of two epiphytic orchids with distinct leaf thickness.

Methods

Tissue morphology and photosynthetic mode (C₃ and/or CAM) were examined in leaves, pseudobulbs and roots of a thick-leaved (Cattleya walkeriana) and a thin-leaved (Oncidium ‘Aloha’) epiphytic orchid. Morphological features were studied comparing the drought-induced physiological responses observed in each organ after 30 d of either drought or well-watered treatments.

Key Results

Cattleya walkeriana, which is considered a constitutive CAM orchid, displayed a clear drought-induced up-regulation of CAM in its thick leaves but not in its non-leaf organs (pseudobulbs and roots). The set of morphological traits of Cattleya leaves suggested the drought-inducible CAM up-regulation as a possible mechanism of increasing water-use efficiency and carbon economy. Conversely, although belonging to an orchid genus classically considered as performing C₃ photosynthesis, Oncidium ‘Aloha’ under drought seemed to express facultative CAM in its roots and pseudobulbs but not in its leaves, indicating that such photosynthetic responses might compensate for the lack of capacity to perform CAM in its thin leaves. Morphological features of Oncidium leaves also indicated lower efficiency in preventing water and CO₂ losses, while aerenchyma ducts connecting pseudobulbs and leaves suggested a compartmentalized mechanism of nighttime carboxylation via phosphoenolpyruvate carboxylase (PEPC) (pseudobulbs) and daytime carboxylation via Rubisco (leaves) in drought-exposed Oncidium plants.

Conclusions

Water availability modulated CAM expression in an organ-compartmented manner in both orchids studied. As distinct regions of the same orchid could perform different photosynthetic pathways and variable degrees of CAM expression depending on the water availability, more attention should be addressed to this in future studies concerning the abundance of CAM plants.     

Striga hermonthica reduces photosynthesis in sorghum: the importance of stomatal limitations and a potential role for ABA?

We report the effects of the root hemiparasite Striga hermonthica (Del.) Benth. on the growth and photosynthesis of two cultivars of sorghum: CSH-1, a susceptible variety, and Ochuti, which shows some tolerance to S. hermonthica in the field. Within 4 d of parasite attachment to the host roots, infected plants of both cultivars were significantly shorter than uninfected controls. At 55 d, infected plants of both cultivars had significantly less shoot and root biomass, and significantly smaller leaf areas than uninfected controls. The dry weight of S. hermonthica attached to host roots was insufficient at this stage to explain the decreased growth in terms of a competing sink for carbon and nitrogen. Leaf chlorophyll and nitrogen per unit area were greater in infected plants of both cultivars compared with control plants. However, whereas photosynthesis and transpiration in young leaves of infected CSH-1 plants declined with time when compared with controls, the rates in infected Ochuti plants were similar to those in uninfected controls throughout the time course of observation. In both cultivars, a strong correlation was observed between the rate of photosynthesis and stomatal conductance during photosynthetic induction, but infection resulted in a much slower induction than in controls. In CSH-1 plants, both steady-state photosynthesis and stomatal conductance were lower than in controls, whereas in leaves of Ochuti steady-state photosynthesis and stomatal conductance eventually reached the same values as in the control leaves. Results from AlC_i analysis and also from determination of ¹³C isotope discrimination were consistent with a stomatal limitation to photosynthesis in the leaves of Striga-infected plants. The concentration of the plant growth regulator abscisic acid (ABA) was measured in the xylem sap of infected CSH-1 plants only, and was found to be twice that of uninfected plants. A possible role of ABA in determining host response to infection by S. hermonthica is discussed.     

Intra-specific variability and the competition–colonisation trade-off: coexistence, abundance and stability patterns

Intra-specific variability often produces an overlap between species distributions of individual performances which can influence competition relations and community dynamics. We analysed a two-species competition–colonisation model of vegetation with intra-specific variability in juvenile growth. On each patch colonised by both species, the winner was the juvenile with higher individual growth. Intra-specific variability disproportionately favoured the more fecund species because the tail of its distribution represented more individuals. In some cases, this process could even lead to a reversal of competition hierarchy and exclusion of the species with higher mean juvenile performance. In the space of species 2 mean growth and fecundity traits, the combinations of traits allowing coexistence with species 1 appeared close to an ideal trade-off curve. Along this curve, species 2 and species 1 coexisted at similar abundance. The balance of relative abundances diminished with the distance of species 2 from this curve. For a given level of relative species performances, coexistence stability increased continuously as species differentiation increased. In contrast to classical models that exhibit abrupt changes of equilibrium community properties when species traits vary, our model displayed continuous changes of these properties in relation to the balance of life traits within and among species. Intra-specific variability allows flexible patterns of community dynamics and could explain discrepancies between observations and classical theories.     

Clonal plants: beyond the patterns—ecological and evolutionary dynamics of asexual reproduction

More than 25 years have passed since publication of the first comprehensive multi-authored landmark volume on the population biology and evolution of clonal organisms (Jackson et al. 1985). Since then, no less than eight symposium volumes or special issues have appeared in scientific journals reporting on advances in the field of clonal plant research, indicating that the study of clonal organisms has remained an important topic in ecological research. The three most recent overviews were published in special issues of this journal (Stuefer et al. 2000; Tolvanen et al. 2004; Sammul et al. 2008), and these are now supplemented with a fourth special issue of Evolutionary Ecology. The articles published here reflect some of the most important contributions to a workshop on clonal plant biology held in Leuven (Belgium) in July 2009 and they illustrate some major advances that have been made over the last few years. In the following paragraphs, we first summarize some representative contributions to the current issue, and second, we put forward some personal ideas about promising and underexplored research lines in clonal plant research.     

Inefficient nitrogen resorption in genets of the actinorhizal nitrogen fixing shrubComptonia peregrina: physiological ineptitude or evolutionary tradeoff?

Nutrient resorption was measured in an actinorhizal nitrogen-fixing shrub,Comptonia peregrina, for five years in the understory of a deciduous oak forest in Rhode Island, USA. Mean resorption of nitrogen was extremely inefficient (11%) compared to most deciduous species (50%+), yet resorption of phosphorus was efficient (53%) and comparable to other species. Of the seven additional nutrients studied, only copper (6%) and zinc (10%) were resorbed from senescing leaves. Resorption of nitrogen (5%–20%) and phosphorus (40%–71%) varied significantly among years. Copper was resorbed from leaves in three years and accreted into leaves in two years. Five-year resorption means differed among individual genets by as much as a factor of 2.5 for nitrogen, and 1.3 for phosphorus. Resorption of nitrogen, copper, and zinc were highly correlated, yet resorption of phosphorus remained autonomous from other nutrients. The ecophysiological tradeoffs inComptonia which have resulted in the cooccurence of actinorhizal nitrogen fixation, inefficient nitrogen resorption, and efficient phosphorus resorption suggest that plant nutrient status does have an impact on resorption efficiency and that the evolution of nutrient conservation strategies is nutrient-specific.     

Mayfly emergence along an oligotrophic Dinaric karst hydrosystem: spatial and temporal patterns,and species–environment relationship

Mayfly emergence was studied in the Plitvice Lakes National Park (Croatia) monthly over a 2-year period in four habitats (springs, streams, mountainous rivers, tufa barriers) using monthly collections of emergence traps. A total of 12 mayfly taxa were recorded. Almost half of the collected specimens belonged to the genus Baetis Leach, 1815, which was recorded at every site, but we were unable to distinguish between two included species (B. rhodani and B. cf. nubecularis). Other abundant species were Centroptilum luteolum (Müller, 1776), Alainites muticus (Linnaeus, 1758), Habrophlebia lauta Eaton 1884, Paraleptophlebia submarginata (Stephens, 1835), Serratella ignita (Poda, 1761), Ephemera danica Müller, 1764 and Rhithrogena braaschi Jacob, 1974. The mayfly assemblages at all sites were dominated by species typical of the rhithral zone, but there was a shift in species composition along a longitudinal gradient (from 720 to 390 m a.s.l.) from dominance of eucrenal–epirhithral to metarhithral–hyporhithral elements and finally to appearance of metapotamal and littoral elements. Two environmental factors, maximum water temperature and mean pH, had the highest influence on the mayfly assemblages. Emergence mainly occurred between March and November and was related to the elevated water temperature. Emergence patterns of some species were in accordance with their typical Central European emergence patterns (e.g. S. ignita, H. lauta) while some others showed certain discrepancies (e.g. longer emergence period in Rh. braaschi and P. submarginata, one generation emergence in A. muticus and variable emergence patterns between the sites and between the two studied years in C. luteolum). The current study provides a significant contribution to the knowledge of mayfly ecology in karst freshwater habitats which forms a basis for further investigation and monitoring of mayflies in this area.     

What explains patterns of species richness? The relative importance of climatic‐niche evolution,morphological evolution,and ecological limits in salamanders

A major goal of evolutionary biology and ecology is to understand why species richness varies among clades. Previous studies have suggested that variation in richness among clades might be related to variation in rates of morphological evolution among clades (e.g., body size and shape). Other studies have suggested that richness patterns might be related to variation in rates of climatic‐niche evolution. However, few studies, if any, have tested the relative importance of these variables in explaining patterns of richness among clades. Here, we test their relative importance among major clades of Plethodontidae, the most species‐rich family of salamanders. Earlier studies have suggested that climatic‐niche evolution explains patterns of diversification among plethodontid clades, whereas rates of morphological evolution do not. A subsequent study stated that rates of morphological evolution instead explained patterns of species richness among plethodontid clades (along with “ecological limits” on richness of clades, leading to saturation of clades with species, given limited resources). However, they did not consider climatic‐niche evolution. Using phylogenetic multiple regression, we show that rates of climatic‐niche evolution explain most variation in richness among plethodontid clades, whereas rates of morphological evolution do not. We find little evidence that ecological limits explain patterns of richness among plethodontid clades. We also test whether rates of morphological and climatic‐niche evolution are correlated, and find that they are not. Overall, our results help explain richness patterns in a major amphibian group and provide possibly the first test of the relative importance of climatic niches and morphological evolution in explaining diversity patterns.     

The role of nitrogen in climate change and the impacts of nitrogen–climate interactions in the United States: foreword to thematic issue

Producing food, transportation, and energy for seven billion people has led to large and widespread increases in the use of synthetic nitrogen (N) fertilizers and fossil fuel combustion, resulting in a leakage of N into the environment as various forms of air and water pollution. The global N cycle is more severely altered by human activity than the global carbon (C) cycle, and reactive N dynamics affect all aspects of climate change considerations, including mitigation, adaptation, and impacts. In this special issue of Biogeochemistry, we present a review of the climate–nitrogen interactions based on a technical report for the United States National Climate Assessment presented as individual papers for terrestrial and aquatic ecosystems, agriculture and human health within the US. We provide a brief overview of each of the paper’s main points and conclusions is presented in this foreword summary.     

Excessive nitrogen and phosphorus in European rivers: 2000–2050

Rivers export nutrients to coastal waters. Excess nutrient export may result in harmful algal blooms and hypoxia, affecting biodiversity, fisheries, and recreation. The purpose of this study is to quantify for European rivers (1) the extent to which N and P loads exceed levels that minimize the risk of harmful algal blooms and (2) the relative shares of sources of N and P in rivers. This may help to identify effective management strategies to reduce coastal eutrophication. We focus on 48 rivers in 27 countries of the European Union (EU27). We used the Global Nutrient Export from Watersheds (NEWS) model to analyze nutrient export by rivers and the associated potentials for coastal eutrophication as reflected by Indicator for Coastal Eutrophication Potential (ICEP). In 2000, 38 of the 48 EU rivers indicated in our study had an ICEP > 0, indicating a relatively high potential for harmful algal blooms. These 38 rivers cover 60% of EU27 land area. Between 2000 and 2050 nutrient export by European rivers is projected to decrease. However, by 2050 still 34 EU rivers, covering 48% of the land area, have an ICEP > 0. This indicates that in these scenarios little progress is made in terms of environmental improvement. About one-third of the rivers with ICEP > 0 are N limited, and about two-thirds P limited. In N-limited rivers reducing N loads is a more effective way to reduce the risk for coastal eutrophication than reducing P, and vice versa. For N-limited rivers agriculture or sewage are the dominant sources of nutrients in river water. In P-limited rivers, sewage is found to be the dominant source of P, except for rivers draining into the Atlantic Ocean, where agriculture can also be dominant. A basin-specific approach is needed to effectively reduce N and P loads.     

Symmetries,non-Euclidean metrics,and patterns in a Swift–Hohenberg model of the visual cortex

The aim of this work is to investigate the effect of the shift-twist symmetry on pattern formation processes in the visual cortex. First, we describe a generic set of Riemannian metrics of the feature space of orientation preference that obeys properties of the shift-twist, translation, and reflection symmetries. Second, these metrics are embedded in a modified Swift-Hohenberg model. As a result we get a pattern formation process that resembles the pattern formation process in the visual cortex. We focus on the final stable patterns that are regular and periodic. In a third step we analyze the influences on pattern formation using weakly nonlinear theory and mode analysis. We compare the results of the present approach with earlier models.