Leaf Area, Competition with Grass, and Clover Cultivar: Key Factors to Successful Overwintering and Fast Regrowth of White Clover (Trifolium repens L.) in Spring

The greater sensitivity of white clover (Trifolium repens L.) to low temperature compared with perennial ryegrass (Lolium perenne L.) is a major problem in sustaining the relative contributions to yield of the two species in mixed swards. The objectives of this investigation were to examine the dynamics of leaf development of two white clover cultivars, AberHerald and Grasslands Huia, under field conditions, and to determine the significance of leaf area in winter, and of competition by perennial ryegrass, for the overwintering and regrowth of white clover in spring. Undefoliated white clover plants developed 3·8-6·6 new leaves between late autumn and early spring, and stolon dry matter and total non-structural carbohydrates (TNC) content increased by 262 and by 68% respectively. In contrast, white clover plants that were defoliated frequently during the winter showed a 28% decrease in stolon dry matter and an 82% decrease in the content of TNC. Frequent defoliation in winter caused severe reduction in the rates of emergence of nodes (by 60%) and of buds (by 67%), and the rate of death of nodes and buds increased by a factor of 10, leading to small stolon systems of individual plants in spring. Competition had similar, but weaker, effects to those of winter defoliation, presumably caused by shading of white clover leaves. AberHerald had a higher cumulative leaf emergence (by 22%), a higher mean leaf number (by 23%), a higher stolon DM (by 36%) and a higher TNC content per plant (by 115%) than Grasslands Huia. Results demonstrate the crucial positive role of leaf area during winter, the negative effect of grass competition, and the importance of the clover cultivar, for the overwintering and subsequent spring regrowth of white clove

Differential expression of α- and β-expansin genes in the elongating leaf of Festuca pratensis

Grasses contain a number of genes encoding both α- and β-expansins. These cell wall proteins are predicted to play a role in cell wall modifications, particularly during tissue elongation. We report here on the characterisation of five α- and three vegetative β-expansins expressed in the leaf elongation zone (LEZ) of the forage grass, Festuca pratensis Huds. The expression of the predominant α-expansin (FpExp2) was localised to the vascular tissue, as was the β-expansin FpExpB3. Expression of another β-expansin (FpExpB2) was not localised to vascular tissue but was highly expressed in roots and initiating tillers. This is the first description of vegetative β-expansin gene expression at the organ and tissue level and also the first evidence of differential expression between members of this gene family. In addition, an analysis of both α- and β-expansin expression along the LEZ revealed no correlation with growth rate distribution, whereas we were able to identify a novel xyloglucan endotransglycosylase (FpXET1) whose expression profile closely mimicked leaf growth rate. These data suggest that α- and β-expansin activities in the grass leaf are associated with tissue differentiation, that expansins involved in leaf growth may represent more minor components of the spectrum of expansin genes expressed in this tissue, and that XETs may be useful markers for the analysis of grass leaf growth

Significance of Legumes for the Distribution of Plant Species in Grassland Ecosystems at Different Altitudes in the Alps

The composition of plant species in grassland communities is determined in part by negative and positive interactions among neighbouring plants; such interactions could be dispersal or - in the case of legumes - symbiotic N2 fixation or resource acquisition. We hypothesised that small scale distribution of plant species may be in part affected by (i) the plant's narrow dispersal in the close neighbourhood, (ii) by the N input into the soil through symbiotic N2 fixation by legumes, and (iii) by small scale heterogeneity of N availability in the soil. To address these hypotheses, in four species-rich semi-natural and natural grasslands along an altitudinal gradient from 900 to 2100ma.s.l., the association of species that grew within a radius of 5cm of selected plant species (central plant, CP) was studied. The distribution of the plant species within the plant community was not random and the species composition that grew within a radius of 5cm of a CP varied with altitude. The strength of the association between a CP and its neighbouring species decreased with increasing altitude. The 5cm radius was most dominated by individuals of the same species as the CP, suggesting that narrow dispersal (either through seeds or through vegetative propagates) may be most important for the species composition within the sward. δ15N values of non leguminous species growing near legumes compared to non leguminous species that grew far from legumes indicate that N from symbiotic N2 fixation may be another factor influencing the association of plant species. Effects of small scale heterogeneity in apparent N availability in the soil were detected to be the least important for the species compositio

Variation in acclimation of photosynthesis in Trifolium repens after eight years of exposure to Free Air CO2 Enrichment (FACE)

The initial stimulation of photosynthesis observed on elevation of [CO2] in grasslands has been predicted to be a transient phenomenon constrained by the loss of photosynthetic capacity due to other limitations, notably nutrients and sinks for carbohydrates. Legumes might be expected partially to escape these feedbacks through symbiotic N2 fixation. The Free‐Air Carbon dioxide Enrichment (FACE) experiment at Eschikon, Switzerland, has been the longest running investigation of the effects of open‐air elevation of [CO2] on vegetation. The prediction of a long‐term loss of photosynthetic capacity was tested by analysing photosynthesis in Trifolium repens L. (cv. Milkanova) in the spring and autumn of the eighth, ninth and tenth years of treatment. A high and low N treatment also allowed a test of the significance of exogenous N‐supply in maintaining a stimulation of photosynthetic capacity in the long‐term. Prior work in this Free Air CO2 Enrichment (FACE) experiment has revealed that elevated [CO2] increased both vegetative and reproductive growth of T. repens independent of N treatment. It is shown here that the photosynthetic response of T. repens was also independent of N fertilization under both current ambient and elevated (600 µmol mol-1) [CO2]. There was a strong effect of season on photosynthesis, with light‐saturated rates (Asat) 37% higher in spring than in autumn. Higher Asat in the spring was supported by higher maximum Rubisco carboxylation rates (Vc,max) and maximum rates of electron transport (Jmax) contributing to RuBP regeneration. Elevated [CO2] increased Asat by 37% when averaged across all measurement periods and both N fertilization levels, and decreased stomatal conductance by 25%. In spring, there was no effect of elevated [CO2] on photosynthetic capacity of leaves, but in autumn both Vc,max and Jmax were reduced by approximately 20% in elevated [CO2]. The results show that acclimation of photosynthetic capacity can occur in a nitrogen‐fixing species, in the field where there are no artificial restrictions on sink capacity. However, even with acclimation there was a highly significant increase in photosynthesis at elevated [CO2

Increased abundance of MTD1 and MTD2 mRNAs in nodules of decapitated Medicago truncatula.

To gain insight into the molecular processes occurring in root nodule metabolism after stress, we used a mRNA differential display (DDRT-PCR) approach to identify cDNAs corresponding to genes whose expression is enhanced in nodules of decapitated Medicago truncatula plants. Two full-length cDNAs of plant origin were isolated (MTD1 and MTD2). Sequence analysis revealed that MTD1 is identical to an EST clone (accession number AW559774) expressed in roots of M. truncatula upon infection with Phytophthora medicaginis, while MTD2 is highly homologous to an Arabidopsis thaliana gene (accession number AL133292) coding for a RNA binding-like protein. The two mRNAs started to accumulate in root nodules at 4 h after plant decapitation and reached even higher transcript levels at 24 h from the imposition of the treatment. MTD1 and MTD2 mRNAs were mainly induced in nodules, with very little induction in roots. The abundance of the two transcripts did not change in response to other perturbations known to decrease nitrogenase activity, such as nitrate and Ar/O2 treatments. Our results suggest that MTD1 and MTD2 represent transcripts that accumulate locally in nodules and may be involved in changes in nodule metabolism in response to decapitation

Soil mineral nitrogen availability was unaffected by elevated atmospheric pCO2 in a four year old field experiment (Swiss FACE)

The effect of elevated (60 Pa) atmospheric carbon dioxide partial pressure (pCO2) and N fertilisation on the availability of mineral N and on N transformation in the soil of a Lolium perenne L. monoculture was investigated in the Swiss FACE (Free Air Carbon dioxide Enrichment) experiment. The apparent availability of nitrate and ammonium for plants was estimated during a representative, vegetative re-growth period at weekly intervals from the sorption of the minerals to mixed-bed ion-exchange resin bags at a soil depth of 5 cm. N mineralisation was measured using sequential coring and in situ exposure of soil cores in the top 10 cm of the soil before and after the first cut in spring 1997. High amounts of mineral N were bound to the ion exchange resin during the first week of re-growth. This was probably the combined result of the fertiliser application, the weak demand for N by the newly cut sward and presumably high rates of root decay and exudation after cutting the sward. During the first 2 weeks after the application of fertiliser N at the first cut, there was a dramatic reduction in available N; N remained low during the subsequent weeks of re-growth in all treatments. Overall, nitrate was the predominant form of mineral N that bound to the resin for the duration of the experiment. Apparently, there was always more nitrate than ammonium available to the plants in the high N fertilisation treatment for the whole re-growth period. Apparent N availability was affected significantly by elevated pCO2 only in the first week after the cut; under high N fertilisation, elevated pCO2 increased the amount of mineral N that was apparently available to the plants. Elevated pCO2 did not affect apparent net transformation of N, loss of N or uptake of N by plants. The present data are consistent with earlier results and suggest that the amount of N available to plants from soil resources does not generally increase under elevated atmospheric pCO2. Thus, a possible limiting effect of N on primary production could become more stringent under elevated atmospheric pCO2 as the demand of the plant for N increase