Crosstalk between plant responses to pathogens and herbivores: a view from the outside in

Plants encounter numerous pests and pathogens in the natural environment. An appropriate response to attack by such organisms can lead to tolerance or resistance mechanisms that enable the plant to survive. Many studies concentrate on the signalling pathways that enable plants to recognize and respond to attack, and measure the downstream effect in either biochemical or molecular terms. At the whole plant level, ecologists examine the fitness costs of attack not only for the plant but also over a range of trophic levels. The links between these differing levels of study are beginning to be addressed by the adoption of molecular approaches in more ecologically relevant settings. This review will describe the different approaches used by ecologists and cell biologists in this field and will try to address the question of how we can explore the response to, and consequences, of attack by multiple enemies.

Relationships between plant traits and climate in the Mediterranean region: an analysis based on pollen data

Question: What are the correlations between the degree of drought stress and temperature, and the adoption of specific adaptive strategies by plants in the Mediterranean region? Location: 602 sites across the Mediterranean region. Method: We considered 12 plant morphological and phenological traits, and measured their abundance at the sites as trait scores obtained from pollen percentages. We conducted stepwise regression analyses of trait scores as a function of plant available moisture (α) and winter temperature (MTCO). Results: Patterns in the abundance for the plant traits we considered are clearly determined by α, MTCO or a combination of both. In addition, trends in leaf size, texture, thickness, pubescence and aromatic leaves and other plant level traits such as thorniness and aphylly, vary according to the life form (tree, shrub, forb), the leaf type (broad, needle) and phenology (evergreen, summer-green). Conclusions: Despite conducting this study based on pollen data we have identified ecologically plausible trends in the abundance of traits along climatic gradients. Plant traits other than the usual life form, leaf type and leaf phenology carry strong climatic signals. Generally, combinations of plant traits are more climatically diagnostic than individual traits. The qualitative and quantitative relationships between plant traits and climate parameters established here will help to provide an improved basis for modelling the impact of climate changes on vegetation and form a starting point for a global analysis of pollen-climate relationships

Nutrient sensing and signalling in plants: Potassium and phosphorus

Potassium and phosphorus are important macronutrients for crops but are often deficient in the field. Very little is known about how plants sense fluctuations in K and P and how information about K and P availability is integrated at the whole plant level into physiological and metabolic adaptations. This chapter reviews recent advances in discovering molecular responses of plants to K and P deficiency by microarray experiments. These studies provide us not only with a comprehensive picture of adaptive mechanisms, but also with a large number of transcriptional markers that can be used to identify upstream components of K and P signalling pathways. On the basis of the available information we discuss putative receptors and signals involved in the sensing and integration of K and P status both at the cellular and at the whole plant level. These involve membrane potential, voltage-dependent ion channels, intracellular Ca and pH, and transcription factors, as well as hormones and metabolites for systemic signalling. Genetic screens of reporter lines for transcriptional markers and metabolome analysis of K- and P-deficient plants are likely to further advance our knowledge in this area in the near future.

Granulometric characterization and evaluation of annually banded mid-Holocene estuarine silts, Welsh Severn Estuary (UK): coastal change, sea level and climate

Holocene silts (salt marshes) and highest intertidal-supratidal peats are superbly exposed on a 15 kin coastal transect which reveals two laterally extensive units of annually banded silts (Beds 3, 7) associated with three transgressive-regressive silt-peat cycles (early sixth-early fourth millennium BC). Bed 3 in places is concordantly and gradationally related to peats above and below, but in others transgresses older strata. Bed 7 also grades up into peat, but everywhere overlies a discordance. The banding in Bed 3 at three main and two minor sites was resolved and characterized texturally at high-resolution (2.5/5 mm contiguous slices) using laser granulometry (LS230 with PIDS) and a comprehensive scheme of data-assessment. Most of Bed 3 formed very rapidly, at peak values of several tens of millimetres annually, in accordance with modelled effects of sea-level fluctuations on mature marshes (bed concordant and gradational) and on marshes growing up after coastal erosion and retreat (bed with discordant base). Using data from the modern Severn Estuary, the textural contrast within bands, and its variation between bands, points to a variable but overall milder mid-Holocene climate than today. The inter-annual variability affected marsh dynamics, as shown by the behaviour of the finely divided plant tissues present. Given local calibration, the methodology is applicable to other tidal systems with banded silts in Britain and mainland northwest Europe. (c) 2006 Elsevier Ltd. All rights reserved.

Influence of host plant heterogeneity on the distribution of a birch aphid

1. The spatial and temporal abundance of the aphid Euceraphis betulae was investigated in relation to heterogeneity in host plant ( Betula pendula) vigour and pathogenic stress. The performance of aphids feeding on vigorous and stressed foliage was also examined. 2. The plant stress and plant vigour hypotheses have been suggested as opposing ways in which foliage quality influences herbivore abundance. In many plants, however, vigorous growing foliage co-exists with stressed or damaged foliage. 3. There was a negative correlation between branch growth ( vigour) and branch stress ( leaf chlorosis), with the most vigorous branches displaying little or no stress, and the most stressed branches achieving poor growth. There was a similar negative correlation between vigour and stress at the level of individual trees, which themselves represented a continuum in quality. 4. At the beginning of the season, E. betulae were intermittently more abundant on vigorous branches than on branches destined to become stressed, but aphids became significantly more abundant on stressed branches later in the season, when symptoms of stress became apparent. Similar patterns of aphid abundance were seen on vigorous and stressed trees in the following year. 5. Euceraphis betulae performance was generally enhanced when feeding on naturally stressed B. pendula leaves, but there was some evidence for elevated potential reproduction when feeding on vigorous leaves too. 6. Overall, plant stress probably influences E. betulae distribution more than plant vigour, but the temporal and spatial variability in plant quality suggests that plant vigour could play a role in aphid distribution early in the season.

Plant species and functional group effects on abiotic and microbial soil properties and plant-soil feedback responses in two grasslands

1 Plant species differ in their capacity to influence soil organic matter, soil nutrient availability and the composition of soil microbial communities. Their influences on soil properties result in net positive or negative feedback effects, which influence plant performance and plant community composition. 2 For two grassland systems, one on a sandy soil in the Netherlands and one on a chalk soil in the United Kingdom, we investigated how individual plant species grown in monocultures changed abiotic and biotic soil conditions. Then, we determined feedback effects of these soils to plants of the same or different species. Feedback effects were analysed at the level of plant species and plant taxonomic groups (grasses vs. forbs). 3 In the sandy soils, plant species differed in their effects on soil chemical properties, in particular potassium levels, but PLFA (phospholipid fatty acid) signatures of the soil microbial community did not differ between plant species. The effects of soil chemical properties were even greater when grasses and forbs were compared, especially because potassium levels were lower in grass monocultures. 4 In the chalk soil, there were no effects of plant species on soil chemical properties, but PLFA profiles differed significantly between soils from different monocultures. PLFA profiles differed between species, rather than between grasses and forbs. 5 In the feedback experiment, all plant species in sandy soils grew less vigorously in soils conditioned by grasses than in soils conditioned by forbs. These effects correlated significantly with soil chemical properties. None of the seven plant species showed significant differences between performance in soil conditioned by the same vs. other plant species. 6 In the chalk soil, Sanguisorba minor and in particular Briza media performed best in soil collected from conspecifics, while Bromus erectus performed best in soil from heterospecifics. There was no distinctive pattern between soils collected from forb and grass monocultures, and plant performance could not be related to soil chemical properties or PLFA signatures. 7 Our study shows that mechanisms of plant-soil feedback can depend on plant species, plant taxonomic (or functional) groups and site-specific differences in abiotic and biotic soil properties. Understanding how plant species can influence their rhizosphere, and how other plant species respond to these changes, will greatly enhance our understanding of the functioning and stability of ecosystems.

Quantifying the effect of buffer zones, crop areas and spatial aggregation on the externalities of genetically modified crops at landscape level

The development of genetically modified (GM) crops has led the European Union (EU) to put forward the concept of 'coexistence' to give fanners the freedom to plant both conventional and GM varieties. Should a premium for non-GM varieties emerge in the market, 'contamination' by GM pollen would generate a negative externality to conventional growers. It is therefore important to assess the effect of different 'policy variables'on the magnitude of the externality to identify suitable policies to manage coexistence. In this paper, taking GM herbicide tolerant oilseed rape as a model crop, we start from the model developed in Ceddia et al. [Ceddia, M.G., Bartlett, M., Perrings, C., 2007. Landscape gene flow, coexistence and threshold effect: the case of genetically modified herbicide tolerant oilseed rape (Brassica napus). Ecol. Modell. 205, pp. 169-180] use a Monte Carlo experiment to generate data and then estimate the effect of the number of GM and conventional fields, width of buffer areas and the degree of spatial aggregation (i.e. the 'policy variables') on the magnitude of the externality at the landscape level. To represent realistic conditions in agricultural production, we assume that detection of GM material in conventional produce might occur at the field level (no grain mixing occurs) or at the silos level (where grain mixing from different fields in the landscape occurs). In the former case, the magnitude of the externality will depend on the number of conventional fields with average transgenic presence above a certain threshold. In the latter case, the magnitude of the externality will depend on whether the average transgenic presence across all conventional fields exceeds the threshold. In order to quantify the effect of the relevant' policy variables', we compute the marginal effects and the elasticities. Our results show that when relying on marginal effects to assess the impact of the different 'policy variables', spatial aggregation is far more important when transgenic material is detected at field level, corroborating previous research. However, when elasticity is used, the effectiveness of spatial aggregation in reducing the externality is almost identical whether detection occurs at field level or at silos level. Our results show also that the area planted with GM is the most important 'policy variable' in affecting the externality to conventional growers and that buffer areas on conventional fields are more effective than those on GM fields. The implications of the results for the coexistence policies in the EU are discussed. (C) 2008 Elsevier B.V. All rights reserved.

Species divergence and trait convergence in experimental plant community assembly

Despite decades of research, it remains controversial whether ecological communities converge towards a common structure determined by environmental conditions irrespective of assembly history. Here, we show experimentally that the answer depends on the level of community organization considered. In a 9-year grassland experiment, we manipulated initial plant composition on abandoned arable land and subsequently allowed natural colonization. Initial compositional variation caused plant communities to remain divergent in species identities, even though these same communities converged strongly in species traits. This contrast between species divergence and trait convergence could not be explained by dispersal limitation or community neutrality alone. Our results show that the simultaneous operation of trait-based assembly rules and species-level priority effects drives community assembly, making it both deterministic and historically contingent, but at different levels of community organization.

The biocontrol fungus Pochonia chlamydosporia shows nematode host preference at the infraspecific level

A RAPD-PCR assay was developed and used to test For competitive variability in growth of the nematode biological control fungus Pochonia chlamydosporia. Saprophytic competence in soil with or without tomato plants was examined in three isolates of the fungus: RES 280 (J), originally isolated from potato cyst nematode (PCN) cysts; RES 200 (1) and RES 279 (S), both originally isolated from root knot nematode (RKN) eggs. Viable counts taken at 70 d indicated that I was the best saprophyte followed by S, with J the poorest. RAPD-PCR analysis of colonies from mixed treatments revealed that there was a cumulative effect of adding isolates to the system. This Suggested that the isolates did not interact and that they may occupy separate niches in soil and the rhizosphere. To investigate parasitic ability, soils were seeded with two isolates of the fungus: J and S, singly or in combination. Tomato or potato plants were grown in these soils; free of nematodes, or inoculated with PCN or RKN, and incubated for 77 d. The abundance of the PCN isolate J in PCN cysts was significantly greater than that of the RKN isolate S but in RKN egg masses, S was significantly more abundant than J. RAPD-PCR analysis of colonies from mixed treatments confirmed that J was more abundant than S ill PCN cysts whereas the converse was observed on RKN egg masses. This substantiates the phenomenon of nematode host preference at the infraspecific level of P. chlamydosporia and highlights its relevance for biological control of plant parasitic nematodes.

Response of plant-pollinator communities to fire: changes in diversity, abundance and floral reward structure

Globally, plant-pollinator communities are subject to a diverse array of perturbations and in many temperate and semi-arid systems fire is a dominant structuring force. We present a novel and highly integrated approach, which quantifies, in parallel, the response to fire of pollinator communities, floral communities and floral reward structure. Mt Carmel, Israel is a recognised bee-flower biodiversity hotspot, and using a chronosequence of habitats with differing post-fire ages, we follow the changes in plant-pollinator community organisation from immediately following a burn until full regeneration of vegetation. Initially, fire has a catastrophic effect on these communities, however, recovery is rapid with a peak in diversity of both flowers and bees in the first 2 years post-fire, followed by a steady decline over the next 50 years. The regeneration of floral communities is closely matched by that of their principal pollinators. At the community level we quantify, per unit area of habitat, key parameters of nectar and pollen forage known to be of importance in structuring pollinator communities. Nectar Volume, nectar water content, nectar concentration and the diversity of nectar foraging niches are all greatest immediately following fire with a steady decrease as regeneration proceeds. Temporal changes in energy availability for nectar, pollen, total energy (nectar + pollen) and relative importance of pollen to nectar energy show a similar general decline with site age, however, the pattern is less clear owing to the highly patchy distribution of floral resources. Changes in floral reward structure reflect the general shift from annuals (generally low-reward open access flowers) to perennials (mostly high-reward and restricted access flowers) as post-fire regeneration ensues. The impact of fire on floral communities and their associated rewards have clear implications for pollinator community structure and we discuss this and the role of other disturbance factors on these systems.

Phylogeny and the hierarchical organization of plant diversity

R. H. Whittaker's idea that plant diversity can be divided into a hierarchy of spatial components from alpha at the within-habitat scale through beta for the turnover of species between habitats to gamma along regional gradients implies the underlying existence of alpha, beta, and gamma niches. We explore the hypothesis that the evolution of a, (3, and gamma niches is also hierarchical, with traits that define the a niche being labile, while those defining a and 7 niches are conservative. At the a level we find support for the hypothesis in the lack of close significant phylogenetic relationship between meadow species that have similar a niches. In a second test, a niche overlap based on a variety of traits is compared between congeners and noncongeners in several communities; here, too, there is no evidence of a correlation between a niche and phylogeny. To test whether beta and gamma niches evolve conservatively, we reconstructed the evolution of relevant traits on evolutionary trees for 14 different clades. Tests against null models revealed a number of instances, including some in island radiations, in which habitat (beta niche) and elevational maximum (an aspect of the gamma niche) showed evolutionary conservatism.

Concentration in the ownership of plant variety rights: Some implications for developing countries

This paper explores concentration levels in the ownership of intellectual property rights over plant varieties worldwide. An analysis of data for 30 UPOV member-countries shows a high degree of concentration in the ownership of plant variety rights for six major crops at the national level in the developed world. Much of this concentration has arisen owing to the rapid consolidation of the seed industry through mergers and acquisitions, especially in the 1990s. A high degree of concentration in the ownership of plant variety rights, in combination with recent efforts to strengthen plant variety protection regimes, is likely to have significant effects on the prospects for future innovation in plant breeding and the distribution of market power between companies. For developing countries, concentration in intellectual property right ownership may have important implications for the structure of domestic seed industries and access to protected varieties and associated plant breeding technologies. These implications for developing countries are likely to become apparent in the context of the rapid spread of plant variety protection and access legislation, emerging changes in the international exchange regime for plant material and liberalised investment policies permitting foreign investment in the seeds sector.

Concentration in ownership of plant variety rights: some implications for developing countries

This paper explores concentration levels in the ownership of intellectual property rights over plant varieties worldwide. An analysis of data for 30 UPOV member-countries shows a high degree of concentration in the ownership of plant variety rights for six major crops at the national level in the developed world. Much of this concentration has arisen owing to the rapid consolidation of the seed industry through mergers and acquisitions, especially in the 1990s. A high degree of concentration in the ownership of plant variety rights, in combination with recent efforts to strengthen plant variety protection regimes, is likely to have significant effects on the prospects for future innovation in plant breeding and the distribution of market power between companies. For developing countries, concentration in intellectual property right ownership may have important implications for the structure of domestic seed industries and access to protected varieties and associated plant breeding technologies. These implications for developing countries are likely to become apparent in the context of the rapid spread of plant variety protection and access legislation, emerging changes in the international exchange regime for plant material and liberalised investment policies permitting foreign investment in the seeds sector. (C) 2003 Elsevier Ltd. All rights reserved.

The effects of cattle grazing on plant-pollinator communities in a fragmented Mediterranean landscape

The main aims of this study were to assess grazing impacts on bee communities in fragmented mediterranean shrubland (phrygana) and woodland habitats that also experience frequent wildfires, and to explain the mechanisms by which these impacts occur. Fieldwork was carried out in 1999 and 2000 on Mount Carmel, in northern Israel, a known hot-spot for bee diversity. Habitats with a range of post-burn ages and varying intensities of cattle grazing were surveyed by transect recording, grazing levels, and the diversity and abundance of both flowers and bees were measured. The species richness of both bees and flowers were highest at moderate to high grazing intensities, and path-analysis indicated that the effects of both grazing and fire on bee diversity were mediated mainly through changes in flower diversity, herb flowers being more important than shrubs. The abundance of bees increased with intensified grazing pressure even at the highest levels surveyed. Surprisingly though, changes in bee abundance at high grazing levels were not caused directly by changes in flower cover. The variation in bee abundance may have been due to higher numbers of solitary bees from the family Halictidae in grazed sites, where compacted ground (nesting resource) and composites (forage resource) were abundant. The effects of grazing on plants were clearest in the intermediate-aged sites, where cattle inhibited the growth of some of the dominant shrubs, creating or maintaining more open patches where light-demanding herbs could grow, thus allowing a diverse flora to develop. Overall, bee communities benefit from a relatively high level of grazing in phrygana. Although bee and flower diversity may decrease under very heavy grazing, the present levels of grazing on Mount Carmel appear to have only beneficial effects on the bee community.

Responses of invertebrate trophic level, feeding guild and body size to the management of improved grassland field margins

P>1. Management of lowland mesotrophic grasslands in north-west Europe often makes use of inorganic fertilizers, high stocking densities and silage-based forage systems to maximize productivity. The impact of these practices has resulted in a simplification of the plant community combined with wide-scale declines in the species richness of grassland invertebrates. We aim to identify how field margin management can be used to promote invertebrate diversity across a suite of functionally diverse taxa (beetles, planthoppers, true bugs, butterflies, bumblebees and spiders). 2. Using an information theoretic approach we identify the impacts of management (cattle grazing, cutting and inorganic fertilizer) and plant community composition (forb species richness, grass species richness and sward architecture) on invertebrate species richness and body size. As many of these management practices are common to grassland systems throughout the world, understanding invertebrate responses to them is important for the maintenance of biodiversity. 3. Sward architecture was identified as the primary factor promoting increased species richness of both predatory and phytophagous trophic levels, as well as being positively correlated with mean body size. In all cases phytophagous invertebrate species richness was positively correlated with measures of plant species richness. 4. The direct effects of management practices appear to be comparatively weak, suggesting that their impacts are indirect and mediated though the continuous measures of plant community structure, such as sward architecture or plant species richness. 5. Synthesis and applications. By partitioning field margins from the remainder of the field, economically viable intensive grassland management can be combined with extensive management aimed at promoting native biodiversity. The absence of inorganic fertilizer, combined with a reduction in the intensity of both cutting and grazing regimes, promotes floral species richness and sward architectural complexity. By increasing sward architecture the total biomass of invertebrates also increased (by c. 60% across the range of sward architectural measures seen in this study), increasing food available for higher trophic levels, such as birds and mammals.