Changing precipitation patterns alter plant community dynamics and succession in an ex-arable grassland

1. Changes in the frequency of extreme events, such as droughts, may be one of the most significant impacts of climate change for ecosystems. Models predict more frequent summer droughts in much of England: this paper investigates the impact on different types of plants in an ex-arable grassland community. 2. A long-term experiment simulated increased and decreased summer precipitation. Substantial interannual variation allowed the effects of summer drought to be tested in combination with wet and dry weather in other seasons. This is important, as climate models predict increased winter precipitation. 3. Total cover abundance in early summer increased with increasing water supply in the previous summer; there was no effect of winter precipitation. Productivity is therefore likely to decrease with more frequent summer droughts, with no mitigating effect of wetter winters. 4. The percentage cover of perennial grasses declined during a natural drought in 1995-97; this was exacerbated by the experimental drought treatment and reduced by supplemented rainfall. Simultaneously, short-lived ruderal species increased; this was greatest in drought treatments and least with supplemented rainfall. 4. These trends were subsequently reversed during several years of unusually wet weather, with perennial grasses increasing and short-lived forbs decreasing. This occurred even in experimentally droughted plots, and we propose that it resulted from rapid coverage of gaps during wet autumns and winters. 6. Deep-rooted species generally proved to be more drought resistant, but there were exceptions. 7. We conclude that increased frequency of summer droughts could have serious implications for the establishment and successional development of ex-arable grasslands. Increased winter precipitation would moderate the impact on species composition, but not on productivity.

Factors affecting distribution and mobility of trace elements (Cu, Pb, Zn) in a perennial grapevine (Vitis vinifera L.) in the Champagne region of France

Soil and Vitis vinifera L (coarse and fine roots, leaves, berries) concentration and geochemical partitioning of Cu, Pb and Zn were determined in a contaminated calcareous Champagne plot to assess their mobility and transfer. Accumulation ratios in roots remained low (0.1-0.4 for Cu and Zn, <0.05 for Pb). Differences between elements resulted from vegetation uptake strategy and soil partitioning. Copper, significantly associated with the oxidisable fraction (27.8%), and Zn with the acid soluble fraction (33.3%), could be mobilised by rhizosphere acidification and oxidisation, unlike Pb, essentially contained in the reducible fraction (72.4%). Roots should not be considered as a whole since the more reactive fine roots showed higher accumulation ratios than coarse ones. More sensitive response of fine roots, lack of correlation between chemical extraction results and vegetation concentrations, and very limited translocation to aerial parts showed that fine root concentrations should be used when assessing bioavailability. (C) 2008 Elsevier Ltd. All rights reserved.

Drought impacts on above-belowground interactions: do effects differ between annual and perennial host species?

Root herbivores can have a positive or negative effect on the abundance and/or performance of foliar phytophages. In addition, abiotic factors such as drought can either strengthen or weaken this effect, depending on the system under investigation. One explanation for these varying responses lies in differences in the physiological response of host plants to drought and root herbivores. Here, the impacts of root phytophages on a leaf-mining species feeding on annual and perennial plant species (four Sonchus species) were compared. The responses of plants and leaf-miners to dtought and root herbivore treatments were not related to whether the host plant was an annual or perennial. However, where root feeders did affect foliar phytophage performance, this occurred only under a drought treatment, demonstrating the potential for climatic change to alter the outcome of plant-mediated interactions. (c) 2007 Gessellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.

Interactions between the grasses Phalaris arundinacea, Miscanthus sinensis and Echinochloa crus-galli, and barley and cereal yellow dwarf virusus

The grass species Miscanthus sinensis, Echinochloa crus-galli and Phalaris arundinacea may be useful biomass crops. In glasshouse inoculations with two isolates of Barley yellow dwarf virus (BYDV)-MAV and BYDV-PAV and one of Cereal yellow dwarf virus (CYVD)-RPV , E. crus galli was infected by all three virus isolates, P. arundinacea by BYDV-MAV and CYDV-RPV, but M. sinensis only by BYDV-MAV. All three hosts became very difficult to infect after several weeks’ growth. Symptoms were inconspicuous; dry matter yield losses ranged from c. 20–40%. Aphids acquired all three virus isolates from E. crus-galli, but more efficiently from 5 than 26-week-old plants. Only BYDV-MAV was acquired from P. arundinacea and M. sinensis. Plants of each species and of Avena sativa were grown outdoors between May and July in 1994 and 1995. Young plants of each species were exposed for successive 2-week intervals during the same periods. Vector populations were higher on A. sativa and P. arundinacea than on E. crus-galli and M. sinensis, and more plants of these species became infected. In 1994 only BYDV-MAV was detected. In 1995 BYDV-MAV, BYDV-PAV and CYDV-RPV were all detected; BYDV-MAV was again the virus isolate most frequently found.

Technologies for biological containment of GM and Non-GM crops

International Perspective The development of GM technology continues to expand into increasing numbers of crops and conferred traits. Inevitably, the focus remains on the major field crops of soybean, maize, cotton, oilseed rape and potato with introduced genes conferring herbicide tolerance and/or pest resistance. Although there are comparatively few GM crops that have been commercialised to date, GM versions of 172 plant species have been grown in field trials in 31 countries. European Crops with Containment Issues Of the 20 main crops in the EU there are four for which GM varieties are commercially available (cotton, maize for animal feed and forage, and oilseed rape). Fourteen have GM varieties in field trials (bread wheat, barley, durum wheat, sunflower, oats, potatoes, sugar beet, grapes, alfalfa, olives, field peas, clover, apples, rice) and two have GM varieties still in development (rye, triticale). Many of these crops have hybridisation potential with wild and weedy relatives in the European flora (bread wheat, barley, oilseed rape, durum wheat, oats, sugar beet and grapes), with escapes (sunflower); and all have potential to cross-pollinate fields non-GM crops. Several fodder crops, forestry trees, grasses and ornamentals have varieties in field trials and these too may hybridise with wild relatives in the European flora (alfalfa, clover, lupin, silver birch, sweet chestnut, Norway spruce, Scots pine, poplar, elm, Agrostis canina, A. stolonifera, Festuca arundinacea, Lolium perenne, L. multiflorum, statice and rose). All these crops will require containment strategies to be in place if it is deemed necessary to prevent transgene movement to wild relatives and non-GM crops. Current Containment Strategies A wide variety of GM containment strategies are currently under development, with a particular focus on crops expressing pharmaceutical products. Physical containment in greenhouses and growth rooms is suitable for some crops (tomatoes, lettuce) and for research purposes. Aquatic bioreactors of some non-crop species (algae, moss, and duckweed) expressing pharmaceutical products have been adopted by some biotechnology companies. There are obvious limitations of the scale of physical containment strategies, addressed in part by the development of large underground facilities in the US and Canada. The additional resources required to grow plants underground incurs high costs that in the long term may negate any advantage of GM for commercial productioNatural genetic containment has been adopted by some companies through the selection of either non-food/feed crops (algae, moss, duckweed) as bio-pharming platforms or organisms with no wild relatives present in the local flora (safflower in the Americas). The expression of pharmaceutical products in leafy crops (tobacco, alfalfa, lettuce, spinach) enables growth and harvesting prior to and in the absence of flowering. Transgenically controlled containment strategies range in their approach and degree of development. Plastid transformation is relatively well developed but is not suited to all traits or crops and does not offer complete containment. Male sterility is well developed across a range of plants but has limitations in its application for fruit/seed bearing crops. It has been adopted in some commercial lines of oilseed rape despite not preventing escape via seed. Conditional lethality can be used to prevent flowering or seed development following the application of a chemical inducer, but requires 100% induction of the trait and sufficient application of the inducer to all plants. Equally, inducible expression of the GM trait requires equally stringent application conditions. Such a method will contain the trait but will allow the escape of a non-functioning transgene. Seed lethality (‘terminator’ technology) is the only strategy at present that prevents transgene movement via seed, but due to public opinion against the concept it has never been trialled in the field and is no longer under commercial development. Methods to control flowering and fruit development such as apomixis and cleistogamy will prevent crop-to-wild and wild-to-crop pollination, but in nature both of these strategies are complex and leaky. None of the genes controlling these traits have as yet been identified or characterised and therefore have not been transgenically introduced into crop species. Neither of these strategies will prevent transgene escape via seed and any feral apomicts that form are arguably more likely to become invasives. Transgene mitigation reduces the fitness of initial hybrids and so prevents stable introgression of transgenes into wild populations. However, it does not prevent initial formation of hybrids or spread to non-GM crops. Such strategies could be detrimental to wild populations and have not yet been demonstrated in the field. Similarly, auxotrophy prevents persistence of escapes and hybrids containing the transgene in an uncontrolled environment, but does not prevent transgene movement from the crop. Recoverable block of function, intein trans-splicing and transgene excision all use recombinases to modify the transgene in planta either to induce expression or to prevent it. All require optimal conditions and 100% accuracy to function and none have been tested under field conditions as yet. All will contain the GM trait but all will allow some non-native DNA to escape to wild populations or to non-GM crops. There are particular issues with GM trees and grasses as both are largely undomesticated, wind pollinated and perennial, thus providing many opportunities for hybridisation. Some species of both trees and grass are also capable of vegetative propagation without sexual reproduction. There are additional concerns regarding the weedy nature of many grass species and the long-term stability of GM traits across the life span of trees. Transgene stability and conferred sterility are difficult to trial in trees as most field trials are only conducted during the juvenile phase of tree growth. Bio-pharming of pharmaceutical and industrial compounds in plants Bio-pharming of pharmaceutical and industrial compounds in plants offers an attractive alternative to mammalian-based pharmaceutical and vaccine production. Several plantbased products are already on the market (Prodigene’s avidin, β-glucuronidase, trypsin generated in GM maize; Ventria’s lactoferrin generated in GM rice). Numerous products are in clinical trials (collagen, antibodies against tooth decay and non-Hodgkin’s lymphoma from tobacco; human gastric lipase, therapeutic enzymes, dietary supplements from maize; Hepatitis B and Norwalk virus vaccines from potato; rabies vaccines from spinach; dietary supplements from Arabidopsis). The initial production platforms for plant-based pharmaceuticals were selected from conventional crops, largely because an established knowledge base already existed. Tobacco and other leafy crops such as alfalfa, lettuce and spinach are widely used as leaves can be harvested and no flowering is required. Many of these crops can be grown in contained greenhouses. Potato is also widely used and can also be grown in contained conditions. The introduction of morphological markers may aid in the recognition and traceability of crops expressing pharmaceutical products. Plant cells or plant parts may be transformed and maintained in culture to produce recombinant products in a contained environment. Plant cells in suspension or in vitro, roots, root cells and guttation fluid from leaves may be engineered to secrete proteins that may be harvested in a continuous, non-destructive manner. Most strategies in this category remain developmental and have not been commercially adopted at present. Transient expression produces GM compounds from non-GM plants via the utilisation of bacterial or viral vectors. These vectors introduce the trait into specific tissues of whole plants or plant parts, but do not insert them into the heritable genome. There are some limitations of scale and the field release of such crops will require the regulation of the vector. However, several companies have several transiently expressed products in clinical and pre-clinical trials from crops raised in physical containment.

Divergent evolutionary pattern of starch biosynthetic pathway genes in grasses and dicots

Starch is the most widespread and abundant storage carbohydrate in crops and its production is critical to both crop yield and quality. As regards the starch content in the seeds of crop plants, there are distinct difference between grasses (Poaceae) and dicots. However, few studies have described the evolutionary pattern of genes in the starch biosynthetic pathway in these two groups of plants. In this study, therefore, an attempt was made to compare the evolutionary rate, gene duplication and selective pattern of the key genes involved in this pathway between the two groups, using five grasses and five dicots as materials. The results showed (i) distinct differences in patterns of gene duplication and loss between grasses and dicots; duplication in grasses mainly occurred prior to the divergence of grasses, whereas duplication mostly occurred in individual species within the dicots; there is less gene loss in grasses than in dicots; (ii) a considerably higher evolutionary rate in grasses than in dicots in most gene families analyzed; (iii) evidence of a different selective pattern between grasses and dicots; positive selection may have occurred asymmetrically in grasses in some gene families, e.g. AGPase small subunit. Therefore, we deduced that gene duplication contributes to, and a higher evolutionary rate is associated with, the higher starch content in grasses. In addition, two novel aspects of the evolution of the starch biosynthetic pathway were observed.

Glycosyl transferases in family 61 mediate arabinofuranosyl transfer onto xylan in grasses

Xylan, a hemicellulosic component of the plant cell wall, is one of the most abundant polysaccharides in nature. In contrast to dicots, xylan in grasses is extensively modified by alpha-(1,2)- and alpha-(1,3)-linked arabinofuranose. Despite the importance of grass arabinoxylan in human and animal nutrition and for bioenergy, the enzymes adding the arabinosyl substitutions are unknown. Here we demonstrate that knocking-down glycosyltransferase (GT) 61 expression in wheat endosperm strongly decreases alpha-(1,3)-linked arabinosyl substitution of xylan. Moreover, heterologous expression of wheat and rice GT61s in Arabidopsis leads to arabinosylation of the xylan, and therefore provides gain-of-function evidence for alpha-(1,3)-arabinosyltransferase activity. Thus, GT61 proteins play a key role in arabinoxylan biosynthesis and therefore in the evolutionary divergence of grass cell walls.

Alternatives to Sedum on green roofs: Can broad leaf perennial plants offer better ‘cooling service’?

Green roof plants alter the microclimate of building roofs and may improve roof insulation. They act by providing cooling by shading, but also through transpiration of water through their stomata. However, leaf surfaces can become warmer when plants close the stomata and decrease water loss in response to drying substrate (typically associated with green roofs during summers), also reducing transpirational cooling. By using a range of contrasting plant types (Sedum mix – an industry green roof ‘standard’, Stachys byzantina, Bergenia cordifolia and Hedera hibernica) we tested the hypothesis that plants differ in their ‘cooling potential’. We firstly examined how leaf morphology influenced leaf temperature and how drying substrate altered that response. Secondly, we investigated the relationship between leaf surface temperatures and the air temperatures immediately above the canopies (i.e. potential to provide aerial cooling). Finally we measured how the plant type influenced the substrate temperature below the canopy (i.e. potential for building cooling). In our experiments Stachys outperformed the other species in terms of leaf surface cooling (even in drying substrate, e.g. 5 oC cooler compared with Sedum), substrate cooling beneath its canopy (up to 12 oC) and even - during short intervals over hottest still periods - the air above the canopy (up to 1 oC, when soil moisture was not limited). We suggest that the choice of plant species on green roofs should not be entirely dictated by what survives on the shallow substrates of extensive systems, but consideration should be given to supporting those species providing the greatest eco-system service potential.

The grass-free lawn: management and species choice for optimum ground cover and plant diversity

In Britain, managed grass lawns provide the most traditional and widespread of garden and landscape practices in use today. Grass lawns are coming under increasing challenge as they tend to support a low level of biodiversity and can require substantial additional inputs to maintain. Here we apply a novel approach to the traditional monocultural lawnscape by replacing grasses entirely with clonal perennial forbs. We monitored changes in plant coverage and species composition over a two year period and here we report the results of a study comparing plant origin native, non-native and mixed) and mowing regime. This allows us to assess the viability of this construct as an alternative to traditional grass lawns. Grass-free lawns provided a similar level of plant cover to grass lawns. Both the mowing regime and the combination of species used affected this outcome, with native plant species seen to have the highest survival rates, and mowing at 4cm to produce the greatest amount of ground coverage and plant species diversity within grass-free lawns. Grass-free lawns required over 50% less mowing than a traditionally managed grass lawn. Observations suggest that plant forms that exhibited: a) a relatively fast growth rate, b) a relatively large individual leaf area, and c) an average leaf height substantially above the cut to be applied, were unsuitable for use in grass-free lawns. With an equivalent level of ground coverage to grass lawns, increased plant diversity and a reduced need for mowing, the grass-free lawn can be seen as a species diverse, lower input and potentially highly ornamental alternative to the traditional lawn format.

Physiological and morphological adaptations of herbaceous perennial legumes allow differential access to sources of varyingly soluble phosphate

The aim of this study was to investigate the capacity of three perennial legume species to access sources of varyingly soluble phosphorus (P) and their associated morphological and physiological adaptations. Two Australian native legumes with pasture potential (Cullen australasicum and Kennedia prostrata) and Medicago sativa cv. SARDI 10 were grown in sand under two P levels (6 and 40 µg P g−1) supplied as Ca(H2PO4)2·H2O (Ca-P, highly soluble, used in many fertilizers) or as one of three sparingly soluble forms: Ca10(OH)2(PO4)6 (apatite-P, found in relatively young soils; major constituent of rock phosphate), C6H6O24P6Na12 (inositol-P, the most common form of organic P in soil) and FePO4 (Fe-P, a poorly-available inorganic source of P). All species grew well with soluble P. When 6 µg P g−1 was supplied as sparingly soluble P, plant dry weight (DW) and P uptake were very low for C. australasicum and M. sativa (0.1–0.4 g DW) with the exception of M. sativa supplied with apatite-P (1.5 g). In contrast, K. prostrata grew well with inositol-P (1.0 g) and Fe-P (0.7 g), and even better with apatite-P (1.7 g), similar to that with Ca-P (1.9 g). Phosphorus uptake at 6 µg P g−1 was highly correlated with total root length, total rhizosphere carboxylate content and total rhizosphere acid phosphatase (EC 3.1.3.2) activity. These findings provide strong indications that there are opportunities to utilize local Australian legumes in low P pasture systems to access sparingly soluble soil P and increase perennial legume productivity, diversity and sustainability.

Contrasting responses to drought stress in herbaceous perennial legumes

Background and aims Medicago sativa L. is widely grown in southern Australia, but is poorly adapted to dry, hot summers. This study aimed to identify perennial herbaceous legumes with greater resistance to drought stress and explore their adaptive strategies. Methods Ten herbaceous perennial legume species/accessions were grown in deep pots in a sandy, low-phosphorus field soil in a glasshouse. Drought stress was imposed by ceasing to water. A companion M. sativa plant in each pot minimised differences in leaf area and water consumption among species. Plants were harvested when stomatal conductance of stressed plants decreased to around 10% of well watered plants. Results A range of responses to drought stress were identified, including: reduced shoot growth; leaf curling; thicker pubescence on leaves and stems; an increased root:shoot ratio; an increase, decrease or no change in root distribution with depth; reductions in specific leaf area or leaf water potential; and osmotic adjustment. The suite of changes differed substantially among species and, less so, among accessions. Conclusions The inter- and intra-specific variability of responses to drought-stress in the plants examined suggests a wide range of strategies are available in perennial legumes to cope with drying conditions, and these could be harnessed in breeding/selection programs.

Variation in seedling growth of 11 perennial legumes in response to phosphorus supply

Phosphorus (P) deficiency is a major problem for Australian agriculture. Development of new perennial pasture legumes that acquire or use P more efficiently than the current major perennial pasture legume, lucerne (Medicago sativa L.), is urgent. A glasshouse experiment compared the response of ten perennial herbaceous legume species to a series of P supplies ranging from 0 to 384 µg g−1 soil, with lucerne as the control. Under low-P conditions, several legumes produced more biomass than lucerne. Four species (Lotononis bainesii Baker, Kennedia prorepens F.Muell, K. prostrata R.Br, Bituminaria bituminosa (L.) C.H.Stirt) achieved maximum growth at 12 µg P g−1 soil, while other species required 24 µg P g−1. In most tested legumes, biomass production was reduced when P supply was ≥192 µg g−1, due to P toxicity, while L. bainesii and K. prorepens showed reduced biomass when P was ≥24 µg g−1 and K. prostrata at ≥48 µg P g−1 soil. B. bituminosa and Glycine canescens F.J.Herm required less soil P to achieve 0.5 g dry mass than the other species did. Lucerne performed poorly with low P supply and our results suggest that some novel perennial legumes may perform better on low-P soils.

Variation in morphological and physiological parameters in herbaceous perennial legumes in response to phosphorus supply

Change in morphological and physiological parameters in response to phosphorus (P) supply was studied in 11 perennial herbaceous legume species, six Australian native (Lotus australis, Cullen australasicum, Kennedia prorepens, K. prostrata, Glycine canescens, C. tenax) and five exotic species (Medicago sativa, Lotononis bainesii, Bituminaria bituminosa var albomarginata, Lotus corniculatus, Macroptilium bracteatum). We aimed to identify mechanisms for P acquisition from soil. Plants were grown in sterilised washed river sand; eight levels of P as KH2PO4 ranging from 0 to 384 μg P g−1 soil were applied. Plant growth under low-P conditions strongly correlated with physiological P-use efficiency and/or P-uptake efficiency. Taking all species together, at 6 μg P g−1 soil there was a good correlation between P uptake and both root surface area and total root length. All species had higher amounts of carboxylates in the rhizosphere under a low level of P application. Six of the 11 species increased the fraction of rhizosphere citrate in response to low P, which was accompanied by a reduction in malonate, except L. corniculatus. In addition, species showed different plasticity in response to P-application levels and different strategies in response to P deficiency. Our results show that many of the 11 species have prospects for low-input agroecosystems based on their high P-uptake and P-use efficiency.