Sugar signaling in root responses to low phosphorus availability

Over the last decade, major advances have been made in our understanding of how plants sense, signal, and respond to soil phosphorus (P) availability (Amtmann et al., 2006; White and Hammond, 2008; Nilsson et al., 2010; Yang and Finnegan, 2010; Vance, 2010; George et al., 2011). Previously, we have reviewed the potential for shoot-derived carbohydrate signals to initiate acclimatory responses in roots to low P availability. In this context, these carbohydrates act as systemic plant growth regulators (Hammond and White, 2008). Photosynthate is transported primarily to sink tissues as Suc via the phloem. Under P starvation, plants accumulate sugars and starch in their leaves. Increased loading of Suc to the phloem under P starvation primarily functions to relocate carbon resources to the roots, which increases their size relative to the shoot (Hermans et al., 2006). The translocation of sugars via the phloem also has the potential to initiate sugar signaling cascades that alter the expression of genes involved plant responses to low P availability. These include optimizing root biochemistry to acquire soil P, through increased expression and activity of inorganic phosphate (Pi) transporters, the secretion of acid phosphatases and organic acids to release P from the soil, and the optimization of internal P use (Hammond and White, 2008). Here, we provide an Update to the field of plant signaling responses to low P availability and the interactions with sugar signaling components. Advances in the P signaling pathways and the roles of hormones in signaling plant responses to low P availability are also reviewed, and where possible their interactions with potential sugar signaling pathways.

Determinants and measures of export performance – comprehensive literature review

As one of the key indicators of the firm’s ability to leverage successfully its resources and capabilities in the international context, export performance has been one of the most extensively studied phenomena. A plethora of studies have been conducted pertaining to provide better understanding of the factors (firm- or environment-specific) and behaviours (e.g., export strategy) that make exporting a successful venture. Following a comprehensive literature review undertaking in this study the current state of the export performance literature could be summarisedas (i) methodologically fragmented in that there is a variety of analytical and methodological approaches, (ii) conceptually diverse, a large number of determinants have been identified as having direct or indirect influence on the firm’s export performance, and a large number of indicators have been used to conceptualise and operationalise the export performance measures, and (iii) inconclusive, the studies have produced inconsistent results of the impact of different determinants on export performance.

Bringing export performance metrics into Annual Reports: the APEV scale and the PERFEX scorecard

Annual company reports rarely distinguish between domestic and export market performance and even more rarely provide information about annual indicators of a specific export venture's performance. In this study, the authors develop and test a new measure for assessing the annual performance of an export venture (the APEV scale). The new measure comprises five dimensions: (1) annual export venture financial performance, (2) annual export venture strategic performance, (3) annual export venture achievement, (4) contribution of the export venture to annual exporting operations, and (5) satisfaction with annual export venture overall performance. The authors use the APEV scale to generate a scorecard of performance in exporting (the PERFEX scorecard) to assess export performance at the corporate level while comparatively evaluating all export ventures of the firm. Both the scale and the scorecard could help disclose export venture performance and could be useful instruments for annual planning, management, monitoring, and improvement of exporting programs.

The RELQUAL scale: a measure of relationship quality in export market ventures

In this article, the authors develop a new measurement scale (the RELQUAL scale) to assess the degree of relationship quality between the exporting firm and the importer. Relationship quality is presented as a high-order concept. Findings reveal that a better quality of the relationship results in a greater (1) amount of information sharing, (2) communication quality, (3) long-term orientation, as well as (4) satisfaction with the relationship. The four multi-item scales show strong evidence of reliability as well as convergent, discriminant and nomological validity in a sample of British exporters. Findings also reveal that relationship quality is positively and significantly associated with export performance. Suggestions for applying the measure in future research are presented.

The STEP scale: a measure of short-term export performance improvement

This article is a direct response to a recent observation in the literature that managers appear to be short-term oriented in their assessment of the performance of an export venture (Madsen 1998). On the basis of a cross-national survey of exporting firms, the authors present a three-dimensional scale for assessing managerial judgment of short-term export performance (i.e., the STEP scale). The three dimensions are (1) satisfaction with short-term performance improvement, (2) short-term exporting intensity improvement, and (3) expected short-term performance improvement. The scale presents evidence of reliability as well as convergent, discriminant, and nomological validity, and it reveals factorial similarity and factorial equivalence across both samples. The authors outline managerial and public policy implications that stem from the scale and identify avenues for further export marketing research.

Characterising phosphorus and nitrate inputs to a rural river using high-frequency concentration-flow relationships

The total reactive phosphorus (TRP) and nitrate concentrations of the River Enborne, southern England, were monitored at hourly interval between January 2010 and December 2011. The relationships between these high-frequency nutrient concentration signals and flow were used to infer changes in nutrient source and dynamics through the annual cycle and each individual storm event, by studying hysteresis patterns. TRP concentrations exhibited strong dilution patterns with increasing flow, and predominantly clockwise hysteresis through storm events. Despite the Enborne catchment being relatively rural for southern England, TRP inputs were dominated by constant, non-rain-related inputs from sewage treatment works (STW) for the majority of the year, producing the highest phosphorus concentrations through the spring–summer growing season. At higher river flows, the majority of the TRP load was derived from within-channel remobilisation of phosphorus from the bed sediment, much of which was also derived from STW inputs. Therefore, future phosphorus mitigation measures should focus on STW improvements. Agricultural diffuse TRP inputs were only evident during storms in the May of each year, probably relating to manure application to land. The nitrate concentration–flow relationship produced a series of dilution curves, indicating major inputs from groundwater and to a lesser extent STW. Significant diffuse agricultural inputs with anticlockwise hysteresis trajectories were observed during the first major storms of the winter period. The simultaneous investigation of high-frequency time series data, concentration–flow relationships and hysteresis behaviour through multiple storms for both phosphorus and nitrate offers a simple and innovative approach for providing new insights into nutrient sources and dynamics.

How well can we model stream phosphorus concentrations in agricultural catchments?

Mechanistic catchment-scale phosphorus models appear to perform poorly where diffuse sources dominate. We investigate the reasons for this for one model, INCA-P, testing model output against 18 months of daily data in a small Scottish catchment. We examine key model processes and provide recommendations for model improvement and simplification. Improvements to the particulate phosphorus simulation are especially needed. The model evaluation procedure is then generalised to provide a checklist for identifying why model performance may be poor or unreliable, incorporating calibration, data, structural and conceptual challenges. There needs to be greater recognition that current models struggle to produce positive Nash–Sutcliffe statistics in agricultural catchments when evaluated against daily data. Phosphorus modelling is difficult, but models are not as useless as this might suggest. We found a combination of correlation coefficients, bias, a comparison of distributions and a visual assessment of time series a better means of identifying realistic simulations.

Palaeoenvironmental changes during the Valanginian: new insights from variations in phosphorus contents and bulk- and clay mineralogies in the western Tethys

Paleoenvironmental and paleoclimatic changes during the Valanginian carbon isotopic excursion (CIE) have been investigated in the western Tethys. For this purpose, bulk-rock and clay mineralogies, as well as phosphorus (P) contents were evaluated in a selection of five sections located in the Vocontian Basin (Angles, SE France; Alvier, E Switzerland; Malleval, E France), and the Lombardian Basin (Capriolo, N Italy; Breggia, S Switzerland). Within the CIE interval, bulk-rock and clay mineralogies are inferred to reflect mostly climate change. The onset of the CIE (Busnardoites campylotoxus ammonite Zone) is characterized by higher detrital index (DI: sum of the detrital minerals divided by calcite contents) values and the presence of kaolinite in their clay-mineral assemblages. In the late Valanginian (from the Saynoceras verrucosum Zone up to the end of the Valanginian), the samples show relatively variable DI and lower values or the absence of kaolinite. The variation in the mineralogical composition is interpreted as reflecting a change from a climate characterized by optimal weathering conditions associated with an increase in terrigenous input on the southern European margin during the CIE towards an overall unstable climate associated with drier conditions in the late Valanginian. This is contrasted by a dissymmetry (proximal vs distal) along the studied transect, the northern Tethyan margin being more sensitive to changes in continental input compared to the distal environments. P accumulation rates (PAR) present similar features. In the Vocontian basin, P content variations are associated with changes in terrigenous influx, whereas in the Lombardian basin (i.e. Capriolo and Breggia), PAR values are less well correlated. This is mainly because the deeper part of the Tethys was less sensitive to changes in continental inputs. The onset of the CIE (top of the B. campylotoxus Zone) records a general increase in PAR suggesting an increase in marine nutrient levels. This is linked to higher continental weathering rates and the enhanced influx of nutrients into the ocean. In the period corresponding to the shift itself, P contents show a dissymmetry between the Vocontian and Lombardian basins (proximal vs distal). For the sections of Malleval, Alvier and Angles, a decrease in P concentrations associated to a decrease in detrital input is observed. In Capriolo and Breggia, PAR show maximum values during the plateau, indicating a more complex interaction between different P sources. The time interval including the top of S. verrucosum Zone up to the end of the Valanginian is characterized by variable PAR values, suggesting variable nutrient influxes. These changes are in agreement with an evolution towards seasonally contrasted conditions in the late Valanginian.

Voluntary export restraints between Britain and Japan: the case of the UK car market (1971–2002)

The rise in international markets of new, productive Japanese car manufacturers provoked intense world competition, which created serious doubts about the economic sustainability of an industry mostly dominated until the 1970s by European and North-American multinational companies. Ultimately, this crisis provoked a deep transformation of the industry, with consequences that had a permanent impact on European companies in the sector. American and later European manufacturers were successful in lobbying governments to provide protection. Using a rich source of data from the UK, I show that the ‘new trade policy’, voluntary export restraint (VER), placed on Japanese exports of new cars from 1977 to December 1999, was binding. This case study illustrates the strategies used by Japanese manufacturers to gain access to the European market through the UK market via strategic alliances and later through transplant production, against which continental European nation states were unable to fully insulate themselves. It is also shown that the policy had a profound effect on the nature of Japanese products, as Japanese firms responded to the quantity restraints by radically altering the product characteristics of their automobiles and shifting towards larger autos and new goods, to maximise their profits subject to the binding constraint.

Strategic trade policy, competition, and welfare: the case of voluntary export restraints between Britain and Japan (1971–2002)

I evaluate the voluntary export restraint placed on Japanese automobile exports from 1977 to 1999 by the UK. I show that the policy failed to assist the British domestic car industry. Instead, UK-based US multi-nationals and Japanese manufacturers were the primary beneficiaries, at a substantial cost to UK consumers. Whilst there are a number of caveats, the policy was on balance damaging to the UK economy in welfare terms.

The forms of bioavailability of phosphorus in integrated vertical flow constructed wetland with earthworms and different substrates

A sequential extraction method was utilized to analyze seven forms of P in an integrated vertical-flow constructed wetland (IVFCW) containing earthworms and different substrates. The aluminum-bound P (Al-P) content was found to be lower, and the occluded P (Oc-P) content was higher in the IVFCW. The addition of earthworms into the influent chamber of IVFCW increased the exchange P (Ex-P), iron-bound P (Fe-P), calcium bound P (Ca-P), Oc-P, detritus-bound (De-P) and organic P (Org-P) content in the influent chamber, and also enhanced P content uptake by wetland plants. A significantly positive correlation between P content of above-ground wetland plants and the Ex-P, Fe-P, Oc-P and Org-P content in the rhizosphere was found (P < 0.05), which indicated that the Ex-P, Fe-P, Oc-P and Org-P could be bio-available P. The Ex-P, Fe-P, De-P, Oc-P and Ca-P content of the influent chamber was higher where the substrate contained a mixture of Qing sand and river sand rather than only river sand. Also the IVFCW with earthworms and both Qing sand and river sand had a higher removal efficiency of P, which was related to higher P content uptake by wetland plants and P retained in IVFCW. These findings suggest that addition of earthworms in IVFCW increases the bioavailable P content, resulting in enhanced P content uptake by wetland plants.

Phosphorus fertilisation and large legume species affect jarrah forest restoration after bauxite mining

Re-establishing nutrient-cycling is often a key goal of mine-site restoration. This goal can be achieved by applying fertilisers (particularly P) in combination with seeding N-fixing legumes. However, the effect of this strategy on other key restoration goals such as the establishment and growth of non-leguminous species has received little attention. We investigated the effects of P-application rates either singly, or in combination with seeding seven large understorey legume species, on jarrah forest restoration after bauxite mining. Five years after P application and seeding, legume species richness, density and cover were higher in the legume-seeded treatment. However, the increased establishment of legumes did not lead to increased soil N. Increasing P-application rates from 0 to 80 kg P ha−1 did not affect legume species richness, but significantly reduced legume density and increased legume cover: cover was maximal (∼50%) where 80 kg P ha−1 had been applied with large legume seeds. Increasing P-application had no effect on species richness of non-legume species, but increased the density of weeds and native ephemerals. Cover of non-legume species decreased with increasing P-application rates and was lower in plots where large legumes had been seeded compared with non-seeded plots. There was a significant legume × P interaction on weed and ephemeral density: at 80 kg P ha−1 the decline in density of these groups was greatest where legumes were seeded. In addition, the decline in cover for non-legume species with increasing P was greatest when legumes were seeded. Applying 20 kg P ha−1 significantly increased tree growth compared with tree growth in unfertilised plots, but growth was not increased further at 80 kg ha−1 and tree growth was not affected by seeding large legumes. Taken together, these data indicate that 80 kg ha−1 P-fertiliser in combination with (seeding) large legumes maximised vegetation cover at five years but could be suboptimal for re-establishing a jarrah forest community that, like unmined forest, contains a diverse community of slow-growing re-sprouter species. The species richness and cover of non-legume understorey species, especially the resprouters, was highest in plots that received either 0 or 20 kg ha−1 P and where large legumes had not been seeded. Therefore, our findings suggest that moderation of P-fertiliser and legumes could be the best strategy to fulfil the multiple restoration goals of establishing vegetation cover, while at the same time maximising tree growth and species richness of restored forest.

Ecto- and arbuscular mycorrhizal symbiosis can induce tolerance to toxic pulses of phosphorus in jarrah (Eucalyptus marginata) seedlings

In common with many plants native to low P soils, jarrah (Eucalyptus marginata) develops toxicity symptoms upon exposure to elevated phosphorus (P). Jarrah plants can establish arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) associations, along with a non-colonizing symbiosis described recently. AM colonization is known to influence the pattern of expression of genes required for P uptake of host plants and our aim was to investigate this phenomenon in relation to P sensitivity. Therefore, we examined the effect on hosts of the presence of AM and ECM fungi in combination with toxic pulses of P and assessed possible correlations between the induced tolerance and the shoot P concentration. The P transport dynamics of AM (Rhizophagus irregularis and Scutellospora calospora), ECM (Scleroderma sp.), non-colonizing symbiosis (Austroboletus occidentalis), dual mycorrhizal (R. irregularis and Scleroderma sp.), and non-mycorrhizal (NM) seedlings were monitored following two pulses of P. The ECM and A. occidentalis associations significantly enhanced the shoot P content of jarrah plants growing under P-deficient conditions. In addition, S. calospora, A. occidentalis, and Scleroderma sp. all stimulated plant growth significantly. All inoculated plants had significantly lower phytotoxicity symptoms compared to NM controls 7 days after addition of an elevated P dose (30 mg P kg−1 soil). Following exposure to toxicity-inducing levels of P, the shoot P concentration was significantly lower in R. irregularis-inoculated and dually inoculated plants compared to NM controls. Although all inoculated plants had reduced toxicity symptoms and there was a positive linear relationship between rank and shoot P concentration, the protective effect was not necessarily explained by the type of fungal association or the extent of mycorrhizal colonization.

Moderating mycorrhizas: arbuscular mycorrhizas modify rhizosphere chemistry and maintain plant phosphorus status within narrow boundaries

Pastures often experience a pulse of phosphorus (P) when fertilized. We examined the role of arbuscular mycorrhizal fungi (AMF) in the uptake of P from a pulse. Five legumes (Kennedia prostrata, Cullen australasicum, Bituminaria bituminosa, Medicago sativa and Trifolium subterraneum) were grown in a moderate P, sterilized field soil, either with (+AMF) or without (−AMF) addition of unsterilized field soil. After 9–10 weeks, half the pots received 15 mg P kg−1 of soil. One week later, we measured: shoot and root dry weights; percentage of root length colonized by AMF; plant P, nitrogen and manganese (Mn) concentrations; and rhizosphere carboxylates, pH and plant-available P. The P pulse raised root P concentration by a similar amount in uncolonized and colonized plants, but shoot P concentration increased by 143% in uncolonized plants and 53% in colonized plants. Inoculation with AMF decreased the amount of rhizosphere carboxylates by 52%, raised rhizosphere pH by ∼0.2–0.7 pH units and lowered shoot Mn concentration by 38%. We conclude that AMF are not simply a means for plants to enhance P uptake when P is limiting, but also act to maintain shoot P within narrow boundaries and can affect nutrient uptake through their influence on rhizosphere chemistry.

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.