Plant-symbiotic fungi as chemical engineers: multi-genome analysis of the Clavicipitaceae reveals dynamics of alkaloid Loci

The fungal family Clavicipitaceae includes plant symbionts and parasites that produce several psychoactive and bioprotective alkaloids. The family includes grass symbionts in the epichloae clade (Epichloë and Neotyphodium species), which are extraordinarily diverse both in their host interactions and in their alkaloid profiles. Epichloae produce alkaloids of four distinct classes, all of which deter insects, and some—including the infamous ergot alkaloids—have potent effects on mammals. The exceptional chemotypic diversity of the epichloae may relate to their broad range of host interactions, whereby some are pathogenic and contagious, others are mutualistic and vertically transmitted (seed-borne), and still others vary in pathogenic or mutualistic behavior. We profiled the alkaloids and sequenced the genomes of 10 epichloae, three ergot fungi (Claviceps species), a morning-glory symbiont (Periglandula ipomoeae), and a bamboo pathogen (Aciculosporium take), and compared the gene clusters for four classes of alkaloids. Results indicated a strong tendency for alkaloid loci to have conserved cores that specify the skeleton structures and peripheral genes that determine chemical variations that are known to affect their pharmacological specificities. Generally, gene locations in cluster peripheries positioned them near to transposon-derived, AT-rich repeat blocks, which were probably involved in gene losses, duplications, and neofunctionalizations. The alkaloid loci in the epichloae had unusual structures riddled with large, complex, and dynamic repeat blocks. This feature was not reflective of overall differences in repeat contents in the genomes, nor was it characteristic of most other specialized metabolism loci. The organization and dynamics of alkaloid loci and abundant repeat blocks in the epichloae suggested that these fungi are under selection for alkaloid diversification. We suggest that such selection is related to the variable life histories of the epichloae, their protective roles as symbionts, and their associations with the highly speciose and ecologically diverse cool-season grasses.

Genome dynamics explain the evolution of flowering time CCT domain gene families in the Poaceae

Numerous CCT domain genes are known to control flowering in plants. They belong to the CONSTANS-like (COL) and PREUDORESPONSE REGULATOR (PRR) gene families, which in addition to a CCT domain possess B-box or response-regulator domains, respectively. Ghd7 is the most recently identified COL gene to have a proven role in the control of flowering time in the Poaceae. However, as it lacks B-box domains, its inclusion within the COL gene family, technically, is incorrect. Here, we show Ghd7 belongs to a larger family of previously uncharacterized Poaceae genes which possess just a single CCT domain, termed here CCT MOTIF FAMILY (CMF) genes. We molecularly describe the CMF (and related COL and PRR) gene families in four sequenced Poaceae species, as well as in the draft genome assembly of barley (Hordeum vulgare). Genetic mapping of the ten barley CMF genes identified, as well as twelve previously unmapped HvCOL and HvPRR genes, finds the majority map to colinear positions relative to their Poaceae orthologues. Combined inter-/intra-species comparative and phylogenetic analysis of CMF, COL and PRR gene families indicates they evolved prior to the monocot/dicot divergence ~200 mya, with Poaceae CMF evolution described as the interplay between whole genome duplication in the ancestral cereal, and subsequent clade-specific mutation, deletion and duplication events. Given the proven role of CMF genes in the modulation of cereals flowering, the molecular, phylogenetic and comparative analysis of the Poaceae CMF, COL and PRR gene families presented here provides the foundation from which functional investigation can be undertaken.

20th century intraseasonal Asian monsoon dynamics viewed from Isomap

The Asian summer monsoon is a high dimensional and highly nonlinear phenomenon involving considerable moisture transport towards land from the ocean, and is critical for the whole region. We have used daily ECMWF reanalysis (ERA-40) sea-level pressure (SLP) anomalies to the seasonal cycle, over the region 50-145°E, 20°S-35°N to study the nonlinearity of the Asian monsoon using Isomap. We have focused on the two-dimensional embedding of the SLP anomalies for ease of interpretation. Unlike the unimodality obtained from tests performed in empirical orthogonal function space, the probability density function, within the two-dimensional Isomap space, turns out to be bimodal. But a clustering procedure applied to the SLP data reveals support for three clusters, which are identified using a three-component bivariate Gaussian mixture model. The modes are found to appear similar to active and break phases of the monsoon over South Asia in addition to a third phase, which shows active conditions over the Western North Pacific. Using the low-level wind field anomalies the active phase over South Asia is found to be characterised by a strengthening and an eastward extension of the Somali jet whereas during the break phase the Somali jet is weakened near southern India, while the monsoon trough in northern India also weakens. Interpretation is aided using the APHRODITE gridded land precipitation product for monsoon Asia. The effect of large-scale seasonal mean monsoon and lower boundary forcing, in the form of ENSO, is also investigated and discussed. The outcome here is that ENSO is shown to perturb the intraseasonal regimes, in agreement with conceptual ideas.

Late Quaternary hydrological dynamics in the Middle Kalahari: forcing and feedbacks

The Kalahari region has become a major source of Quaternary palaeoenvironmental data derived primarily from the analysis of geomorphological proxies of environmental change. One suite of data, from palaeolacustrine landforms, has recently provided a new record of major hydrological changes in the last 150 ka [Burrough, S. L., Thomas, D. S. G., Bailey, R. M., 2009. Mega-Lake in the Kalahari: A Late Pleistocene record of the Palaeolake Makgadikgadi system. Quaternary Science Reviews, in press.]. Here we present an improved analysis of the drivers and feedbacks of lake level change, utilising information from three main sources: data from the lake system itself, from analyses of other late Quaternary records within the region and from climate modelling. Simulations using the Hadley Centre coupled climate model, HadCM3, suggest that once triggered, the lake body was large enough to potentially affect both local and regional climates. Surface waters and their interactions with the climate are therefore an important component of environmental dynamics during the late Quaternary. Through its capacity to couple Middle Kalahari environments to distant forcing mechanisms and to itself force environmental change, we demonstrate that the existence or absence of megalake Makgadikgadi adds a new level of complexity to the interpretations of environmental proxy records in southern Africa's summer rainfall zone.

Effects of plant pathogens on population dynamics and community composition in grassland ecosystems: two case studies

Grassland ecosystems comprise a major portion of the earth’s terrestrial surface, ranging from high-input cultivated monocultures or simple species mixtures to relatively unmanaged but dynamic systems. Plant pathogens are a component of these systems with their impact dependent on many interacting factors, including grassland species population dynamics and community composition, the topics covered in this paper. Plant pathogens are affected by these interactions and also act reciprocally by modifying their nature. We review these features of disease in grasslands and then introduce the 150-year long-term Park Grass Experiment (PGE) at Rothamsted Research in the UK. We then consider in detail two plant-pathogen systems present in the PGE, Tragopogon pratensis-Puccinia hysterium and Holcus lanata-Puccinia coronata. These two systems have very different life history characteristics: the first, a biennial member of the Asteraceae infected by its host-specific, systemic rust; the second, a perennial grass infected by a host-non-specific rust. We illustrate how observational, experimental and modelling studies can contribute to a better understanding of population dynamics, competitive interactions and evolutionary outcomes. With Tragopogon pratensis-Puccinia hysterium, characterised as an “outbreak” species in the PGE, we show that pathogen-induced mortality is unlikely to be involved in host population regulation; and that the presence of even a short-lived seed-bank can affect the qualitative outcomes of the host-pathogen dynamics. With Holcus lanata-Puccinia coronata, we show how nutrient conditions can affect adaptation in terms of host defence mechanisms, and that co-existence of competing species affected by a common generalist pathogen is unlikely.

The role of soil microbes in the global carbon cycle: tracking the below-ground microbial processing of plant-derived carbon for manipulating carbon dynamics in agricultural systems.

It is well known that atmospheric concentrations of carbon dioxide (CO2) (and other greenhouse gases) have increased markedly as a result of human activity since the industrial revolution. It is perhaps less appreciated that natural and managed soils are an important source and sink for atmospheric CO2 and that, primarily as a result of the activities of soil microorganisms, there is a soil-derived respiratory flux of CO2 to the atmosphere that overshadows by tenfold the annual CO2 flux from fossil fuel emissions. Therefore small changes in the soil carbon cycle could have large impacts on atmospheric CO2 concentrations. Here we discuss the role of soil microbes in the global carbon cycle and review the main methods that have been used to identify the microorganisms responsible for the processing of plant photosynthetic carbon inputs to soil. We discuss whether application of these techniques can provide the information required to underpin the management of agro-ecosystems for carbon sequestration and increased agricultural sustainability. We conclude that, although crucial in enabling the identification of plant-derived carbon-utilising microbes, current technologies lack the high-throughput ability to quantitatively apportion carbon use by phylogentic groups and its use efficiency and destination within the microbial metabolome. It is this information that is required to inform rational manipulation of the plant–soil system to favour organisms or physiologies most important for promoting soil carbon storage in agricultural soil.

The population dynamics of disease on short and long time-scales

Observational evidence is scarce concerning the distribution of plant pathogen population sizes or densities as a function of time-scale or spatial scale. For wild pathosystems we can only get indirect evidence from evolutionary patterns and the consequences of biological invasions.We have little or no evidence bearing on extermination of hosts by pathogens, or successful escape of a host from a pathogen. Evidence over the last couple of centuries from crops suggest that the abundance of particular pathogens in the spectrum affecting a given host can vary hugely on decadal timescales. However, this may be an artefact of domestication and intensive cultivation. Host-pathogen dynamics can be formulated mathematically fairly easily–for example as SIR-type differential equation or difference equation models, and this has been the (successful) focus of recent work in crops. “Long-term” is then discussed in terms of the time taken to relax from a perturbation to the asymptotic state. However, both host and pathogen dynamics are driven by environmental factors as well as their mutual interactions, and both host and pathogen co-evolve, and evolve in response to external factors. We have virtually no information about the importance and natural role of higher trophic levels (hyperpathogens) and competitors, but they could also induce long-scale fluctuations in the abundance of pathogens on particular hosts. In wild pathosystems the host distribution cannot be modelled as either a uniform density or even a uniform distribution of fields (which could then be treated as individuals). Patterns of short term density-dependence and the detail of host distribution are therefore critical to long-term dynamics. Host density distributions are not usually scale-free, but are rarely uniform or clearly structured on a single scale. In a (multiply structured) metapopulation with coevolution and external disturbances it could well be the case that the time required to attain equilibrium (if it exists) based on conditions stable over a specified time-scale is longer than that time-scale. Alternatively, local equilibria may be reached fairly rapidly following perturbations but the meta-population equilibrium be attained very slowly. In either case, meta-stability on various time-scales is a more relevant than equilibrium concepts in explaining observed patterns.

Computational fluid dynamics modelling of the air movement in an environmental test chamber with a respiring manikin

In recent years, computational fluid dynamics (CFD) has been widely used as a method of simulating airflow and addressing indoor environment problems. The complexity of airflows within the indoor environment would make experimental investigation difficult to undertake and also imposes significant challenges on turbulence modelling for flow prediction. This research examines through CFD visualization how air is distributed within a room. Measurements of air temperature and air velocity have been performed at a number of points in an environmental test chamber with a human occupant. To complement the experimental results, CFD simulations were carried out and the results enabled detailed analysis and visualization of spatial distribution of airflow patterns and the effect of different parameters to be predicted. The results demonstrate the complexity of modelling human exhalation within a ventilated enclosure and shed some light into how to achieve more realistic predictions of the airflow within an occupied enclosure.

Looking in the wrong place for healthcare improvements: a system dynamics study of an accident and emergency department

Accident and Emergency (A&E) units provide a route for patients requiring urgent admission to acute hospitals. Public concern over long waiting times for admissions motivated this study, whose aim is to explore the factors which contribute to such delays. The paper discusses the formulation and calibration of a system dynamics model of the interaction of demand pattern, A&E resource deployment, other hospital processes and bed numbers; and the outputs of policy analysis runs of the model which vary a number of the key parameters. Two significant findings have policy implications. One is that while some delays to patients are unavoidable, reductions can be achieved by selective augmentation of resources within, and relating to, the A&E unit. The second is that reductions in bed numbers do not increase waiting times for emergency admissions, their effect instead being to increase sharply the number of cancellations of admissions for elective surgery. This suggests that basing A&E policy solely on any single criterion will merely succeed in transferring the effects of a resource deficit to a different patient group.

Should system dynamics be described as a 'hard' or 'deterministic' systems approach?

This paper explores the criticism that system dynamics is a ‘hard’ or ‘deterministic’ systems approach. This criticism is seen to have four interpretations and each is addressed from the perspectives of social theory and systems science. Firstly, system dynamics is shown to offer not prophecies but Popperian predictions. Secondly, it is shown to involve the view that system structure only partially, not fully, determines human behaviour. Thirdly, the field's assumptions are shown not to constitute a grand content theory—though its structural theory and its attachment to the notion of causality in social systems are acknowledged. Finally, system dynamics is shown to be significantly different from systems engineering. The paper concludes that such confusions have arisen partially because of limited communication at the theoretical level from within the system dynamics community but also because of imperfect command of the available literature on the part of external commentators. Improved communication on theoretical issues is encouraged, though it is observed that system dynamics will continue to justify its assumptions primarily from the point of view of practical problem solving. The answer to the question in the paper's title is therefore: on balance, no.

Social theory and system dynamics practice

This paper explores the social theories implicit in system dynamics (SD) practice. Groupings of SD practice are observed in different parts of a framework for studying social theories. Most are seen to be located within `functionalist sociology'. To account for the remainder, two new forms of practice are discussed, each related to a different paradigm. Three competing conclusions are then offered: 1. The implicit assumption that SD is grounded in functionalist sociology is correct and should be made explicit. 2. Forrester's ideas operate at the level of method not social theory so SD, though not wedded to a particular social theoretic paradigm, can be re-crafted for use within different paradigms. 3. SD is consistent with social theories which dissolve the individual/society divide by taking a dialectical, or feedback, stance. It can therefore bring a formal modelling approach to the `agency/structure' debate within social theory and so bring SD into the heart of social science. The last conclusion is strongly recommended.

The greater whole: towards a synthesis of system dynamics and soft systems methodology

This paper makes a theoretical case for using these two systems approaches together. The theoretical and methodological assumptions of system dynamics (SD) and soft system methodology (SSM) are briefly described and a partial critique is presented. SSM generates and represents diverse perspectives on a problem situation and addresses the socio-political elements of an intervention. However, it is weak in ensuring `dynamic coherence'. consistency between the intuitive behaviour resulting from proposed changes and behaviour deduced from ideas on causal structure. Conversely, SD examines causal structures and dynamic behaviours. However, whilst emphasising the need for a clear issue focus, it has little theory for generating and representing diverse issues. Also, there is no theory for facilitating sensitivity to socio-political elements. A synthesis of the two called ‘Holon Dynamics' is proposed. After an SSM intervention, a second stage continues the socio-political analysis and also operates within a new perspective which values dynamic coherence of the mental construct - the holon - which is capable of expressing the proposed changes. A model of this holon is constructed using SD and the changes are thus rendered `systemically desirable' in the additional sense that dynamic consistency has been confirmed. The paper closes with reflections on the proposal and the need for theoretical consistency when mixing tools is emphasised.

Re-fitting Etton: space, time and material culture within a causewayed enclosure in Cambridgeshire

This paper considers the dynamics of deposition around and across the causewayed enclosure at Etton, Cambridgeshire. As a result of detailed re-analysis (particularly refitting) of the pottery and flint assemblages from the site, it proved possible to shed new light both on the temporality of occupation and the character of deposition there. Certain aspects of our work challenge previous interpretations of the site, and of causewayed enclosures in general; but, just as importantly, others confirm materially what has previously been suggested. The quantities of material deposited at Etton reveal that the enclosure was occupied only very intermittently and certainly less regularly than other contemporary sites in the region. The spatial distribution of material suggests that the enclosure ditch lay open for the entirety of the monument's life, but that acts of deposition generally focused on a specific part of the monument at any one time. As well as enhancing our knowledge of one particular causewayed enclosure, it is hoped that this paper – in combination with our earlier analysis of the pit site at Kilverstone – makes clear the potential that detailed material analysis has to offer in relation to our understanding of the temporality of occupation on prehistoric sites in general.

Analysis of the regional pattern of sea level change due to ocean dynamics and density changes for 1993-2099 in observations and CMIP5 AOGCMs

Predictions of twenty-first century sea level change show strong regional variation. Regional sea level change observed by satellite altimetry since 1993 is also not spatially homogenous. By comparison with historical and pre-industrial control simulations using the atmosphere–ocean general circulation models (AOGCMs) of the CMIP5 project, we conclude that the observed pattern is generally dominated by unforced (internal generated) variability, although some regions, especially in the Southern Ocean, may already show an externally forced response. Simulated unforced variability cannot explain the observed trends in the tropical Pacific, but we suggest that this is due to inadequate simulation of variability by CMIP5 AOGCMs, rather than evidence of anthropogenic change. We apply the method of pattern scaling to projections of sea level change and show that it gives accurate estimates of future local sea level change in response to anthropogenic forcing as simulated by the AOGCMs under RCP scenarios, implying that the pattern will remain stable in future decades. We note, however, that use of a single integration to evaluate the performance of the pattern-scaling method tends to exaggerate its accuracy. We find that ocean volume mean temperature is generally a better predictor than global mean surface temperature of the magnitude of sea level change, and that the pattern is very similar under the different RCPs for a given model. We determine that the forced signal will be detectable above the noise of unforced internal variability within the next decade globally and may already be detectable in the tropical Atlantic.