Hidden branches: developments in root system architecture.

The root system is fundamentally important for plant growth and survival because of its role in water and nutrient uptake. Therefore, plants rely on modulation of root system architecture (RSA) to respond to a changing soil environment. Although RSA is a highly plastic trait and varies both between and among species, the basic root system morphology and its plasticity are controlled by inherent genetic factors. These mediate the modification of RSA, mostly at the level of root branching, in response to a suite of biotic and abiotic factors. Recent progress in the understanding of the molecular basis of these responses suggests that they largely feed through hormone homeostasis and signaling pathways. Novel factors implicated in the regulation of RSA in response to the myriad endogenous and exogenous signals are also increasingly isolated through alternative approaches such as quantitative trait locus analysis.

Predicting root defence against herbivores during succession

P>1. Root herbivores and pathogens interfere with basic below-ground plant function, and can thereby affect plant fitness and spatial and temporal patterns in natural plant communities. However, there has been little development of concepts and theories on below-ground plant defence, a deficit that is in contrast to the abundance of theorizing for above-ground plant parts.2. A review of the past 10 years of research on below-ground plant-herbivore interactions has revealed that, similar to above-ground tissues, root defences can be expressed constitutively or induced upon herbivore attack, and can be classified into direct and indirect traits, tolerance, and escape. Indeed, it has been shown that roots tolerate herbivory by outgrowing or re-growing lost tissues, or resist it by producing secondary metabolites that are toxic to herbivores or attract natural enemies of herbivores.3. We propose that, similar to above-ground plant-herbivore theories, the partition of abiotic and biotic factors over ecological succession can serve as the basis for predicting investment in defence strategies below-ground.4. Investigation of herbivore pressure and root responses along primary and secondary successional gradients suggests that: (i) roots are often fast growing, thinner and softer in early compared to later succession. (ii) Insect and nematode herbivore pressure increases until mid-succession and later decreases. (iii) Mycorrhizal abundance increases with succession, and the composition of fungal species changes through succession, often shifting from arbuscular mycorrhizae to ecto-mycorrhizae.5. Based on these findings, and on classical (above-ground) plant defence theory, we suggest the following set of testable hypotheses for below-ground plant defence: (i) During succession, early plants invest most of their resources in growth and less in defences (associated with a general lack of herbivores and pathogens, and with limited availability of resources in the system), therefore relying more on re-growth (tolerance) strategies. (ii) During mid-succession, a buildup of herbivore pressure facilitates replacement by plant species that exhibit greater direct and indirect defence strategies. (iii) Constitutive and inducible levels of defences may trade-off, and early successional plants should rely more on induction of defences after herbivore attack, whereas late successional plants will increasingly rely on constitutively produced levels of physical and chemical defence. (iv) Successional changes in microbial associations have consequences for root defence by improving plant nutrition and defence expression as well as directly competing for root space; however, toxic or impenetrable root defences may also limit association with root symbionts, and so may constrain the expression of root defence.

Does wheat genetically modified for disease resistance affect root-colonizing pseudomonads and arbuscular mycorrhizal fungi?

This study aimed to evaluate the impact of genetically modified (GM) wheat with introduced pm3b mildew resistance transgene, on two types of root-colonizing microorganisms, namely pseudomonads and arbuscular mycorrhizal fungi (AMF). Our investigations were carried out in field trials over three field seasons and at two locations. Serial dilution in selective King's B medium and microscopy were used to assess the abundance of cultivable pseudomonads and AMF, respectively. We developed a denaturing gradient gel electrophoresis (DGGE) method to characterize the diversity of the pqqC gene, which is involved in Pseudomonas phosphate solubilization. A major result was that in the first field season Pseudomonas abundances and diversity on roots of GM pm3b lines, but also on non-GM sister lines were different from those of the parental lines and conventional wheat cultivars. This indicates a strong effect of the procedures by which these plants were created, as GM and sister lines were generated via tissue cultures and propagated in the greenhouse. Moreover, Pseudomonas population sizes and DGGE profiles varied considerably between individual GM lines with different genomic locations of the pm3b transgene. At individual time points, differences in Pseudomonas and AMF accumulation between GM and control lines were detected, but they were not consistent and much less pronounced than differences detected between young and old plants, different conventional wheat cultivars or at different locations and field seasons. Thus, we conclude that impacts of GM wheat on plant-beneficial root-colonizing microorganisms are minor and not of ecological importance. The cultivation-independent pqqC-DGGE approach proved to be a useful tool for monitoring the dynamics of Pseudomonas populations in a wheat field and even sensitive enough for detecting population responses to altered plant physiology.

Interactions between magnesium, calcium, and aluminum on soybean root elongation

Alleviation of Al rhizotoxicity by Ca and Mg can differ among species and genotypes. Root elongation of soybean [Glycine max (L.) Merr.] line N93-S-179 and cvs. Young and Ransom exposed to varying concentrations of Al, Ca and Mg were compared in two experiments using a vertically split root system. Roots extending from a surface compartment with limed soil grew for 12 days into a subsurface compartment with nutrient solution treatments maintained at pH 4.6 with either 0 or 15 µmol L-1 Al. Calcium and Mg concentrations in treatments ranging from 0 to 20 mmol L-1. Although an adequate supply of Mg was provided in the surface soil compartment for soybean top growth, an inclusion of Mg was necessary in the subsurface solutions to promote root elongation in both the presence and absence of Al. In the absence of Al in the subsurface solution, tap root length increased by 74 % and lateral root length tripled when Mg in the solutions was increased from 0 to either 2 or 10 mmol L-1. In the presence of 15 µmol L-1 Al, additions of 2 or 10 mmol L-1 Mg increased tap root length fourfold and lateral root length by a factor of 65. This high efficacy of Mg may have masked differences in Al tolerance between genotypes N93 and Young. Magnesium was more effective than Ca in alleviating Al rhizotoxicity, and its ameliorative properties could not be accounted for by estimated electrostatic changes in root membrane potential and Al3+ activity at the root surface. The physiological mechanisms of Mg alleviation of Al injury in roots, however, are not known.

Inducing effect of skeletal muscle extracts on the appearance of calbindin-immunoreactive dorsal root ganglion cells in culture.

Calbindin D-28k is a calcium-binding protein which is not expressed by dorsal root ganglion cells cultured from 6-day-old (E6) chick embryos. When soluble muscle extracts from embryos at E11, E18 or chickens 2 weeks after hatching were added immediately after seeding, dorsal root ganglia cells grown at E6 displayed neuronal subpopulations expressing calbindin immunoreactivity with time; the effect of muscle extract on the percentage of calbindin-immunoreactive dorsal root ganglia cells followed a dose-response curve. When muscle extract was added to cultures after a 3 day delay, the percentage of calbindin-expressing neurons was unchanged. The effect produced by muscle extract and, to a lesser degree, skin extract on the appearance of calbindin-positive neurons was not reproduced by brain or liver extracts while all four exerted a trophic action on cultured neurons. Hence it is assumed that muscle extract contains a factor which produces an inductive effect on the initiation of calbindin-expression by uncommitted subpopulations of sensory neurons rather than a trophic influence on the selective survival of covertly committed neuronal subpopulations. The fact that muscle extract promoted calbindin expression by dorsal root ganglia cells in neuron-enriched as well as in mixed dorsal root ganglion cell cultures indicates that the factor would act directly on sensory neurons rather than indirectly through mediation of non-neuronal cells. Since the active muscular factor was non-dialysable, heat-inactivated, trypsin-sensitive and retained by molecular filters with a cut-off of 30 K, this factor is probably a protein.

Brain-derived neurotrophic factor-like immunoreactivity is localized mainly in small sensory neurons of rat dorsal root ganglia.

Brain-derived neurotrophic factor (BDNF) is a protein capable of supporting the survival and fiber outgrowth of peripheral sensory neurons. It has been argued that histological detection of BDNF has proven difficult because of its low molecular weight and relatively low expression. In the present study we report that rapid removal of dorsal root ganglia (DRG) from the rat, followed by rapid freezing and appropriate fixation with cold acetone, preserves BDNF in situ without altering protein antigenicity. Under these conditions, specific BDNF-like immunoreactivity was detected in DRG both in vivo and in vitro. During DRG development in vivo, BDNF-like immunoreactivity (BDNF-LI) was observed only in a subset of sensory neurons. BDNF-LI was confined to small neurons, after neurons became morphologically distinct on the basis of size. BDNF-L immunoprecipitate was detected only in neuronal cells, and not in satellite or Schwann cells. While in vivo BDNF localization was restricted to small neurons, practically all neurons in DRG cell culture displayed BDNF-LI. Small or large primary afferent neurons exhibited a faint but clear BDNF-LI during the whole life span of cultures. Again, non-neuronal cells were devoid of BDNF-LI. In conclusion, in DRG in vivo, specific BDNF-LI was confined to small B sensory neurons. In contrast, all DRG sensory neurons displayed BDNF-LI in vitro. The finding that BDNF expressed in all DRG neurons in vitro but not in vivo suggests that BDNF expression may be modulated by environmental factors.

The global regulator GacA of Pseudomonas fluorescens CHA0 is required for the suppression of root diseases in dicotyledons but not in Gramineae

Pseudomonas fluorescens strain CHA0 suppresses various plant diseases caused by soil-borne fungi. The pseudomonad produces the antimicrobial metabolites 2,4-diacetylphloroglucinol (Phl), pyoluteorin (Plt) and hydrogen cyanide, which are important for disease suppression, as well as the siderophores pyoverdine (Pvd), salicylic acid (Sal) and pyochelin (Pch). In the current work, a derivative of CHA0 with a mutation in the global regulator gene gacA (GacA−), which is unable to produce Phl, Plt and HCN, failed to protect the dicotyledonous plants cress and cucumber against damping-off caused by Pythium ultimum. In contrast, the GacA− mutant could still protect the Gramineae wheat and maize against damping-off mediated by the same strain of P. ultimum, and wheat against take-all caused by Gaeumannomyces graminis. However, the GacA− mutant overproduced Pch and Pvd. To gain more insight into disease protection afforded by the GacA− mutant, a GacA− Pvd− double mutant (strain CHA496) was constructed by gene replacement. Strain CHA496 overproduced Pch and Sal compared with CHA0 and protected wheat against P. ultimum and G. graminis, whereas cress and cucumber were not protected. Addition of FeCl3 repressed Pch and Sal production by strain CHA496 in vitro and impaired the protection of wheat in soil microcosms. In conclusion, a functional gacA gene was necessary for the protection of dicotyledons against root diseases, but not for that of Gramineae. Results indicated also that Pch and/or Sal were involved in the ability of the GacA− Pvd− mutant of CHA0 to suppress root diseases in Gramineae.

Root abundance of maize in conventionally-tilled and zero-tilled soils of Argentina

Maize root growth is negatively affected by compacted layers in the surface (e.g. agricultural traffic) and subsoil layers (e.g. claypans). Both kinds of soil mechanical impedances often coexist in maize fields, but the combined effects on root growth have seldom been studied. Soil physical properties and maize root abundance were determined in three different soils of the Rolling Pampa of Argentina, in conventionally-tilled (CT) and zero-tilled (ZT) fields cultivated with maize. In the soil with a light Bt horizon (loamy Typic Argiudoll, Chivilcoy site), induced plough pans were detected in CT plots at a depth of 0-0.12 m through significant increases in bulk density (1.15 to 1.27 Mg m-3) and cone (tip angle of 60 º) penetrometer resistance (7.18 to 9.37 MPa in summer from ZT to CT, respectively). This caused a reduction in maize root abundance of 40-80 % in CT compared to ZT plots below the induced pans. Two of the studied soils had hard-structured Bt horizons (clay pans), but in only one of them (silty clay loam Abruptic Argiudoll, Villa Lía site) the expected penetrometer resistance increases (up to 9 MPa) were observed with depth. In the other clay pan soil (silty clay loam Vertic Argiudoll, Pérez Millán site), penetrometer resistance did not increase with depth but reached 14.5 MPa at 0.075 and 0.2 m depth in CT and ZT plots, respectively. However, maize root abundance was stratified in the first 0.2 m at the Villa Lía and Pérez Millán sites. There, the hard Bt horizons did not represent an absolute but a relative mechanical impedance to maize roots, by the observed root clumping through desiccation cracks.

Level shifts in a panel data based unit root test : an application to the rate of unemployment

Several unit root tests in panel data have recently been proposed. The test developed by Harris and Tzavalis (1999 JoE) performs particularly well when the time dimension is moderate in relation to the cross-section dimension. However, in common with the traditional tests designed for the unidimensional case, it was found to perform poorly when there is a structural break in the time series under the alternative. Here we derive the asymptotic distribution of the test allowing for a shift in the mean, and assess the small sample performance. We apply this new test to show how the hypothesis of (perfect) hysteresis in Spanish unemployment is rejected in favour of the alternative of the natural unemployment rate, when the possibility of a change in the latter is considered.

New evidence of the real interest rate parity for OECD countries using panel unit root tests with breaks

This paper tests for real interest parity (RIRP) among the nineteen major OECD countries over the period 1978:Q2-1998:Q4. The econometric methods applied consist of combining the use of several unit root or stationarity tests designed for panels valid under cross-section dependence and presence of multiple structural breaks. Our results strongly support the fulfilment of the weak version of the RIRP for the studied period once dependence and structural breaks are accounted for.

Effect of soil-spraying time on root-colonization ability of antagonistic Streptomyces griseoviridis

Selostus: Kasvualustan käsittelyajan vaikutus Streptomyces griseoviridis -antagonistin juurten asutuskykyyn

Changes in anatomy and root cell ultrastructure of soybean genotypes under manganese stress

The deleterious effects of both Mn deficiency and excess on the development of plants have been evaluated with regard to aspects of shoot anatomy, ultrastructure and biochemistry, focusing mainly on the manifestation of visual symptoms. However, there is little information in the literature on changes in the root system in response to Mn supply. The objective of this study was to evaluate the effects of Mn doses (0.5, 2.0 and 200.0 μmol L-1) in a nutrient solution on the anatomy of leaves and roots of the Glycine max (L.) cultivars Santa Rosa, IAC-15 and IAC-Foscarin 31. Visual deficiency symptoms were first observed in Santa Rosa and IAC-15, which were also the only cultivars where Mn-toxicity symptoms were observed. Only in IAC-15, a high Mn supply led to root diameter thickening, but without alteration in cells of the bark, epidermis, exodermis and endodermis. The degree of disorganization of the xylem vessels, in particular the metaxylem, differed in the cultivars. Quantity and shape of the palisade parenchyma cells were influenced by both Mn deficiency and toxicity. A reduction in the number of chloroplasts was observed in the three Mn-deficient genotypes. The anatomical alterations in IAC-15 due to nutritional stress were greater, as expressed in extensive root cell cytoplasm disorganization and increased vacuolation at high Mn doses. The degree of changes in the anatomical and ultrastructural organization of roots and leaves of the soybean genotypes studied differed, suggesting the existence of tolerance mechanisms to different intensities of Mn deficiency or excess.

Neuronal expression of the ubiquitin ligase Nedd4-2 in rat dorsal root ganglia: Modulation in the spared nerve injury model of neuropathic pain.

Neuronal hyperexcitability following peripheral nerve lesions may stem from altered activity of voltage-gated sodium channels (VGSCs), which gives rise to allodynia or hyperalgesia. In vitro, the ubiquitin ligase Nedd4-2 is a negative regulator of VGSC α-subunits (Na(v)), in particular Na(v)1.7, a key actor in nociceptor excitability. We therefore studied Nedd4-2 in rat nociceptors, its co-expression with Na(v)1.7 and Na(v)1.8, and its regulation in pathology. Adult rats were submitted to the spared nerve injury (SNI) model of neuropathic pain or injected with complete Freund's adjuvant (CFA), a model of inflammatory pain. L4 dorsal root ganglia (DRG) were analyzed in sham-operated animals, seven days after SNI and 48h after CFA with immunofluorescence and Western blot. We observed Nedd4-2 expression in almost 50% of DRG neurons, mostly small and medium-sized. A preponderant localization is found in the non-peptidergic sub-population. Additionally, 55.7±2.7% and 55.0±3.6% of Nedd4-2-positive cells are co-labeled with Na(v)1.7 and Na(v)1.8 respectively. SNI significantly decreases the proportion of Nedd4-2-positive neurons from 45.9±1.9% to 33.5±0.7% (p<0.01) and the total Nedd4-2 protein to 44%±0.13% of its basal level (p<0.01, n=4 animals in each group, mean±SEM). In contrast, no change in Nedd4-2 was found after peripheral inflammation induced by CFA. These results indicate that Nedd4-2 is present in nociceptive neurons, is downregulated after peripheral nerve injury, and might therefore contribute to the dysregulation of Na(v)s involved in the hyperexcitability associated with peripheral nerve injuries.

Root system distribution of sugar cane as related to nitrogen fertilization, evaluated by two methods: monolith and probes

Few studies on sugar cane have evaluated the root system of the crop, in spite of its importance. This is mainly due to the difficulty of evaluation and high variability of results. The objective of this study was to develop an evaluation method of the cane root system by means of probes so as to evaluate the mass, distribution and metabolically active roots related to N fertilization at planting. For this purpose, an experiment was conducted in an Arenic Kandiustults with medium texture in Jaboticabal/SP, in a randomized block design with four replications and four treatments: control (without N) and 40, 80 and 120 kg ha-1 of N applied in the form of urea in the planting furrow of the cane variety SP81 3250. One week before harvest, a urea-15N solution was applied at the cane stalk base to detect active metabolism in the root system. Trenches of 1.5 m length and 0.6 m depth were opened between two sugar cane rows for root sampling by two methods: monoliths (0.3, 0.2 and 0.15 m wide, deep and long respectively) taken from the trench wall and by probe (internal diameter 0.055 m). For each method, 15 samples per plot were collected. The roots were separated from the soil in a sieve (2 mm mesh), oven-dried (at 65 ºC) and the dry matter was measured. Root sampling by probes resulted in root mass that did not differ from the evaluation in monoliths, indicating that this evaluation method may be used for sugar cane root mass, although neither the root distribution in the soil profile nor the rhizome mass were efficiently evaluated, due to the small sample volume. Nitrogen fertilization at planting did not result in a greater root accumulation in the sugar cane plant, but caused changes in the distribution of the root system in the soil. The absence of N fertilization led to a better root distribution in the soil profile, with 50, 34 and 16 % in the 0-0.2, 0.2-0.4 and 0.4-0.6 m layers, respectively; in the fertilized treatments the roots were concentrated in the surface layer, with on average 70, 17 and 13 % for the same layers. The metabolically active roots were concentrated in the center of the cane stool, amounting to 40 % of the total root mass, regardless of N fertilization (application of 120 kg ha-1 N or without N).

Phosphorus and root distribution and corn growth as related to long-term tillage systems and fertilizer placement

Soil and fertilizer management during cultivation can affect crop productivity and profitability. Long-term experiments are therefore necessary to determine the dynamics of nutrient and root distribution as related to soil profile, as well as the effects on nutrient uptake and crop growth. An 18-year experiment was conducted at the Federal University of Rio Grande do Sul State (UFRGS), in Eldorado do Sul, Brazil, on Rhodic Paleudult soil. Black oat and vetch were planted in the winter and corn in the summer. The soil management methods were conventional, involving no-tillage and strip tillage techniques and broadcast, row-and strip-applied fertilizer placement (triple superphosphate). Available P (Mehlich-1) and root distribution were determined in soil monoliths during the corn grain filling period. Corn shoot dry matter production and P accumulation during the 2006/2007 growing season were determined and the efficiency of P utilization calculated. Regardless of the degree of soil mobilization, P and roots were accumulated in the fertilized zone with time, mainly in the surface layer (0-10 cm). Root distribution followed P distribution for all tillage systems and fertilizer treatments. Under no-tillage, independent of the fertilizer placement, the corn plants developed more roots than in the other tillage systems. Although soil tillage systems and fertilizer treatments affected P and root distribution throughout the soil profile, as well as P absorption and corn growth, the efficiency of P utilization was not affected.