An arbuscular mycorrhizal fungus accelerates decomposition and acquires nitrogen directly from organic material

Arbuscular mycorrhizal fungi (order Glomales), which form mycorrhizal symbioses with two out of three of all plant species, are believed to be obligate biotrophs that are wholly dependent on the plant partner for their carbon supply. It is thought that they possess no degradative capability and that they are unable to decompose complex organic molecules, the form in which most soil nutrients occur. Earlier suggestions that they could exist saprotrophically were based on observation of hyphal proliferation on organic materials. In contrast, other mycorrhizal types have been shown to acquire nitrogen directly from organic sources. Here we show that the arbuscular mycorrhizal symbiosis can both enhance decomposition of and increase nitrogen capture from complex organic material (grass leaves) in soil. Hyphal growth of the fungal partner was increased in the presence of the organic material, independently of the host plant

Root system architecture determines fitness in an Arabidopsis mutant in competition for immobile phosphate ions but not for nitrate ions

Plant root systems often have complex branching patterns. Models indicate that a complex architecture is only required for the acquisition of immobile resources, such as phosphate; mobile ions, notably nitrate, can be effectively taken up by very restricted root systems. We have tested this prediction using the axr4 mutation of Arabidopsis thaliana, the principal phenotypic effect of which is to reduce the number of lateral roots. Arabidopsis thaliana is not a host for mycorrhizal fungi and so acquires all its nutrients through the root system. In both a pot experiment and a field experiment conducted under natural conditions for A. thaliana, we found that only phosphate, and not nitrate, affected the fitness of the mutant relative to the isogenic wild-type line, Columbia. These results confirm model predictions and have implications both for the evolution of complex root systems and for the design of efficient root systems for crops

Plant root proliferation in nitrogen-rich patches confers competitive advantage

Plants respond to environmental heterogeneity, particularly below ground, where spectacular root proliferations in nutrient-rich patches may occur. Such 'foraging' responses apparently maximize nutrient uptake and are now prominent in plant ecological theory. Proliferations in nitrogen-rich patches are difficult to explain adaptively, however. The high mobility of soil nitrate should limit the contribution of proliferation to N capture. Many experiments on isolated plants show only a weak relation between proliferation and N uptake. We show that N capture is associated strongly with proliferation during interspecific competition for finite, locally available, mixed N sources, precisely the conditions under which N becomes available to plants on generally infertile soils. This explains why N-induced root proliferation is an important resource-capture mechanism in N-limited plant communities and suggests that increasing proliferation by crop breeding or genetic manipulation will have a limited impact on N capture by well-fertilized monocultures

Nitrate and phosphate availability and distribution have different effects on root system architecture of Arabidopsis

Plant root systems can respond to nutrient availability and distribution by changing the three-dimensional deployment of their roots: their root system architecture (RSA). We have compared RSA in homogeneous and heterogeneous nitrate and phosphate supply in Arabidopsis. Changes in nitrate and phosphate availability were found to have contrasting effects on primary root length and lateral root density, but similar effects on lateral root length. Relative to shoot dry weight (DW), primary root length decreased with increasing nitrate availability, while it increased with increasing phosphate supply. Lateral root density remained constant across a range of nitrate supplies, but decreased with increasing phosphate supply. In contrast, lateral root elongation was suppressed both by high nitrate and high phosphate supplies. Local supplies of high nitrate or phosphate in a patch also had different effects. Primary root growth was not affected by a high nitrate patch, but growth through a high phosphate patch reduced primary root growth after the root left the patch. A high nitrate patch induced an increase in lateral root density in the patch, whereas lateral root density was unaffected by a high phosphate patch. However, both phosphate- and nitrate-rich patches induced lateral root elongation in the patch and suppressed it outside the patch. This co-ordinated response of lateral roots also occurs in soil-grown plants exposed to a nutrient-rich patch. The auxin-resistant mutants axr1, axr4 and aux1 all showed the wild-type lateral root elongation responses to a nitrate-rich patch, suggesting that auxin is not required for this response

Phosphate availability regulates root system architecture in Arabidopsis

Plant root systems are highly plastic in their development and can adapt their architecture in response to the prevailing environmental conditions. One important parameter is the availability of phosphate, which is highly immobile in soil such that the arrangement of roots within the soil will profoundly affect the ability of the plant to acquire this essential nutrient. Consistent with this, the availability of phosphate was found to have a marked effect on the root system architecture of Arabidopsis. Low phosphate availability favored lateral root growth over primary root growth, through increased lateral root density and length, and reduced primary root growth mediated by reduced cell elongation. The ability of the root system to respond to phosphate availability was found to be independent of sucrose supply and auxin signaling. In contrast, shoot phosphate status was found to influence the root system architecture response to phosphate availability

Rapid turnover of hyphae of mycorrhizal fungi determined by AMS microanalysis of C-14

Processes in the soil remain among the least well-characterized components of the carbon cycle. Arbuscular mycorrhizal (AM) fungi are ubiquitous root symbionts in many terrestrial ecosystems and account for a large fraction of photosynthate in a wide range of ecosystems; they therefore play a key role in the terrestrial carbon cycle. A large part of the fungal mycelium is outside the root ( the extraradical mycelium, ERM) and, because of the dispersed growth pattern and the small diameter of the hyphae (<5 micrometers), exceptionally difficult to study quantitatively. Critically, the longevity of these. ne hyphae has never been measured, although it is assumed to be short. To quantify carbon turnover in these hyphae, we exposed mycorrhizal plants to fossil ("carbon-14 - dead") carbon dioxide and collected samples of ERM hyphae ( up to 116 micrograms) over the following 29 days. Analyses of their carbon-14 content by accelerator mass spectrometry (AMS) showed that most ERM hyphae of AM fungi live, on average, 5 to 6 days. This high turnover rate reveals a large and rapid mycorrhizal pathway of carbon in the soil carbon cycle

Thermal acclimation of leaf and root respiration: an investigation comparing inherently fast- and slow-growing plant species

We investigated the extent to which leaf and root respiration W differ in their response to short- and long-term changes in temperature in several contrasting plant species (herbs, grasses, shrubs and trees) that differ in inherent relative growth rate (RGR, increase in mass per unit starting mass and time). Two experiments were conducted using hydroponically grown plants. In the long-term (LT) acclimation experiment, 16 species were grown at constant 18,23 and 28degreesC. In the short-term (ST) acclimation experiment, 9 of those species were grown at 25/20degreesC (day/night) and then shifted to a 15/10degreesC for 7 days. Short-term Q(10) values (proportional change in R per 10degreesC) and the degree of acclimation to. longer-term changes in temperature were compared. The effect of growth temperature on root and leaf soluble sugar and nitrogen concentrations was examined. Light-saturated photosynthesis (A(sat)) was also measured in the LT acclimation experiment. Our results show that Q(10) values and the degree of acclimation are highly variable amongst species and that roots exhibit lower Q(10) values than leaves over the 15-25degreesC measurement temperature range. Differences in RGR or concentrations of soluble sugars/nitrogen could not account for the inter-specific differences in the Q(10) or degree of acclimation. There were no systematic differences in the ability of roots and leaves to acclimate when plants developed under contrasting temperatures (LT acclimation). However, acclimation was greater in both leaves and roots that developed at the growth temperature (LT acclimation) than in pre-existing leaves and roots shifted from one temperature to another (ST acclimation). The balance between leaf R and A(sat) was maintained in plants grown at different temperatures, regardless of their inherent relative growth rate. We conclude that there is tight coupling between the respiratory acclimation and the temperature under which leaves and roots developed and that acclimation plays an important role in determining the relationship between respiration and photosynthesis

Nitrogen forms affect root structure and water uptake in the hybrid poplar

The study analyses the effects of two different forms of nitrogen fertilisation (nitrate and ammonium) on root structure and water uptake of two hybrid poplar (Populus maximowiczii x P. balsamifera) clones in a field experiment. Water uptake was studied using sap flow gauges on individual proximal roots and coarse root structure was examined by excavating 18 whole-root systems. Finer roots were scanned and analyzed for architecture. Nitrogen forms did not affect coarse-root system development, but had a significant effect on fine-root development. Nitrate-treated trees presented higher fine:coarse root ratios and higher specific root lengths than control or ammonium treated trees. These allocation differences affected the water uptake capacity of the plants as reflected by the higher sapflow rate in the nitrate treatment. The diameter of proximal roots at the tree base predicted well the total root biomass and length. The diameter of smaller lateral roots also predicted the lateral root mass, length, surface area and the number of tips. The effect of nitrogen fertilisation on the fine root structure translated into an effect on the functioning of the fine roots forming a link between form (architecture) and function (water uptake)

Selectivity and functional diversity in arbuscular mycorrhizas of co-occurring fungi and plants from a temperate deciduous woodland

1 The arbuscular mycorrhizal (AM) fungi colonizing plants at a woodland site in North Yorkshire (UK) have been characterized from the roots of five plant species (Rubus fruticosus agg. L., Epilobium angustifolium L., Acer pseudoplatanus L., Ajuga reptans L. and Glechoma hederacea L.), and identified using small-subunit rRNA (SSUrRNA) gene amplification and sequencing. 2 Interactions between five plant species from the site and four co-occurring glomalean fungi were investigated in artificial one-to-one AM symbioses. Three of the fungi were isolated from the site; the fourth was a culture genetically similar to a taxon found at the site. Phosphorus uptake and growth responses were compared with non-mycorrhizal controls. 3 Individual fungi colonized each plant with different spatial distribution and intensity. Some did not colonize at all, indicating incompatibility under the conditions used in the experiments. 4 Glomus hoi consistently occupied a large proportion of root systems and outperformed the other fungi, improving P uptake and enhancing the growth of four out of the five plant species. Only G. hoi colonized and increased P uptake in Acer pseudoplatanus, the host plant with which it associates almost exclusively under field conditions. Colonization of all plant species by Scutellospora dipurpurescens was sparse, and beneficial to only one of the host plants (Teucrium scorodonia). Archaeospora trappei and Glomus sp. UY1225 had variable effects on the host plants, conferring a range of P uptake and growth benefits on Lysimachia nummularia and T. scorodonia, increasing P uptake whilst not affecting biomass in Ajuga reptans and Glechoma hederacea, and failing to form mycorrhizas with A. pseudoplatanus. 5 These experimental mycorrhizas show that root colonization, symbiont compatibility and plant performance vary with each fungus-plant combination, even when the plants and fungi naturally co-exist. 6 We provide evidence of physical and functional selectivity in AM. The small number of described AM fungal species (154) has been ascribed to their supposed lack of host specificity, but if the selectivity we have observed is the general rule, then we may predict that many more, probably hard-to-culture glomalean species await discovery, or that members of species as currently perceived may be physiologically or functionally distinct

Dynamics of hydration water in deuterated purple membranes explored by neutron scattering

The function and dynamics of proteins depend on their direct environment, and much evidence has pointed to a strong coupling between water and protein motions. Recently however, neutron scattering measurements on deuterated and natural-abundance purple membrane (PM), hydrated in H2O and D2O, respectively, revealed that membrane and water motions on the ns–ps time scale are not directly coupled below 260 K (Wood et al. in Proc Natl Acad Sci USA 104:18049–18054, 2007). In the initial study, samples with a high level of hydration were measured. Here, we have measured the dynamics of PM and water separately, at a low-hydration level corresponding to the first layer of hydration water only. As in the case of the higher hydration samples previously studied, the dynamics of PM and water display different temperature dependencies, with a transition in the hydration water at 200 K not triggering a transition in the membrane at the same temperature. Furthermore, neutron diffraction experiments were carried out to monitor the lamellar spacing of a flash-cooled deuterated PM stack hydrated in H2O as a function of temperature. At 200 K, a sudden decrease in lamellar spacing indicated the onset of long-range translational water diffusion in the second hydration layer as has already been observed on flash-cooled natural-abundance PM stacks hydrated in D2O (Weik et al. in J Mol Biol 275:632–634, 2005), excluding thus a notable isotope effect. Our results reinforce the notion that membrane-protein dynamics may be less strongly coupled to hydration water motions than the dynamics of soluble proteins