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Selecting plants with improved root hair growth is a key strategy for improving phosphorus-uptake efficiency in agriculture. While significant inter- and intra-specific variation is reported for root hair length, it is not known whether these phenotypic differences are exhibited under conditions that are known to affect root hair elongation. This work investigates the effect of soil strength, soil water content (SWC) and soil particle size (SPS) on the root hair length of different root hair genotypes of barley. The root hair and rhizosheath development of five root hair genotypes of barley ( Hordeum vulgare L.) was compared in soils with penetrometer resistances ranging from 0.03 to 4.45 MPa (dry bulk densities 1.2-1.7 g cm). A 'short' (SRH) and 'long' root hair (LRH) genotype was selected to further investigate whether differentiation of these genotypes was related to SWC or SPS when grown in washed graded sand. In low-strength soil (<1.43 MPa), root hairs of the LRH genotype were on average 25 % longer than that of the SRH genotype. In high-strength soil, root hair length of the LRH genotype was shorter than that in low-strength soil and did not differ from that of the SRH genotype. Root hairs were shorter in wetter soils or soils with smaller particles, and again SRH and LRH did not differ in hair length. Longer root hairs were generally, but not always, associated with larger rhizosheaths, suggesting that mucilage adhesion was also important. The root hair growth of barley was found to be highly responsive to soil properties and this impacted on the expression of phenotypic differences in root hair length. While root hairs are an important trait for phosphorus acquisition in dense soils, the results highlight the importance of selecting multiple and potentially robust root traits to improve resource acquisition in agricultural systems. [ABSTRACT FROM AUTHOR]

Rhizosheaths are aggregated, sheath-like soils that physically adhere to root surface, and they form on herbaceous plant roots worldwide, especially in semiarid grasslands. Representing a strong root−soil−microbe interaction, the rhizosheaths are expected to have distinct soil organic carbon (SOC) signatures from rhizosphere soils of non-rhizosheath forming plants. However, such signatures remain unclear, which hinders our understanding of root effects on SOC cycling in grasslands. We compared SOC characteristics between rhizosheath and non-rhizosheath soils of eight herbaceous plant species, collected from a semiarid grassland of North China, using solid-state 13C nuclear magnetic resonance spectroscopy and biomarker analyses. We further examined the temporal dynamics of SOC characteristics of rhizosheath soils from early, middle, and late plant growth stages. Compared to non-rhizosheath SOC, rhizosheath SOC had more root inputs of both labile substrates (carbohydrates and free alkanoic acids) and relatively recalcitrant suberin- and lignin-derived compounds. Moreover, the labile inputs provided more substrates for microbial degradation of cutin-derived compounds. These indicators of labile substrate availability increased significantly from the early to late growth stages. Overall, our findings clarify the molecular characteristics of rhizosheath SOC and its temporal dynamics, both of which suggest a critical role of rhizosheath in shaping the rhizosphere microenvironment and regulating SOC cycling. [Display omitted] • Molecular signature of SOC in rhizosheath and non-rhizosheath soils was studied. • More labile compounds increased in rhizosheath soil than non-rhizosheath soil. • Alkyl/ O -alkyl ratio of rhizosheath SOC decreased from early to late growth stages. • Rhizosheaths had more plant inputs and a higher degree of cutin decomposition. [ABSTRACT FROM AUTHOR]

Background and Aims: Wetting-drying cycles are important environmental processes known to enhance aggregation. However, very little attention has been given to drying as a process that transports mucilage to inter-particle contacts where it is deposited and serves as binding glue. The objective of this study was to formulate and test conceptual and mathematical models that describe the role of drying in soil aggregation through transportation and deposition of binding agents. Methods: We used an ESEM to visualize aggregate formation of pair of glass beads. To test our model, we subjected three different sizes of sand to multiple wetting-drying cycles of PGA solution as a mimic of root exudates to form artificial aggregates. Water stable aggregate was determined using wet sieving apparatus. Results: A model to predict aggregate stability in presence of organic matter was developed, where aggregate stability depends on soil texture as well as the strength, density and mass fraction of organic matter, which was confirmed experimentally. The ESEM images emphasize the role of wetting-drying cycles on soil aggregate formation. Conclusions: Our experimental results confirmed the mathematical model predictions as well as the ESEM images on the role of drying in soil aggregation as an agent for transport and deposition of binding agents. [ABSTRACT FROM AUTHOR]

Root hairs are a key trait for improving the acquisition of phosphorus (P) by plants. However, it is not known whether root hairs provide significant advantage for plant growth under combined soil stresses, particularly under conditions that are known to restrict root hair initiation or elongation (e.g. compacted or high-strength soils). To investigate this, the root growth and P uptake of root hair genotypes of barley, Hordeum vulgare L. (i.e. genotypes with and without root hairs), were assessed under combinations of P deficiency and high soil strength. Genotypes with root hairs were found to have an advantage for root penetration into high-strength layers relative to root hairless genotypes. In P-deficient soils, despite a 20% reduction in root hair length under high-strength conditions, genotypes with root hairs were also found to have an advantage for P uptake. However, in fertilized soils, root hairs conferred an advantage for P uptake in low-strength soil but not in high-strength soil. Improved root–soil contact, coupled with an increased supply of P to the root, may decrease the value of root hairs for P acquisition in high-strength, high-P soils. Nevertheless, this work demonstrates that root hairs are a valuable trait for plant growth and nutrient acquisition under combined soil stresses. Selecting plants with superior root hair traits is important for improving P uptake efficiency and hence the sustainability of agricultural systems. [ABSTRACT FROM AUTHOR]

In arid to semi-arid grasslands and savannas, plant growth, population dynamics, and productivity are consistently and strongly limited by soil water and nutrient availability. Adaptive traits of the root systems of grasses in these ecosystems are crucial to their ability to cope with strong water and/or nutrient limitation and the increasing drought stress associated with ecosystem degradation or projected climate change. We studied 18 grass species in semi-arid savanna of the Kalahari region of Botswana to quantify interspecific variation in three important root system traits including root system architecture, rhizosheath thickness and mycorrhizal colonization. Drought-tolerant species and shorter-lived species showed greater rhizosheath thickness and fine root development but lower mycorrhizal colonization compared to later successional climax grasses and those characteristic of wetter sites. In addition, there was a significant positive correlation between root fibrousness index and rhizosheath thickness among species and a weak negative correlation between root fibrousness index and mycorrhizal colonization. These patterns suggest that an extensive fine root system and rhizosheath development may be important complementary traits of grasses coping with drought conditions, the former aiding in the acquisition of water by the grass plant and the latter aiding in water uptake and retention, and reducing water loss in the rhizosphere. Within species, both rhizosheath development and mycorrhizal colonization were significantly greater in a wet year than in a year with below-average precipitation. The observed patterns suggest that the primary benefit of rhizosheath development in African savanna grasses is improved drought tolerance and that it is a plastic trait that can be adjusted annually to changing environmental conditions. The functioning of mycorrhizal symbiosis is likely to be relatively more important in infertile savannas where nutrient limitation is higher relative to water limitation. [ABSTRACT FROM AUTHOR]

Abstract: North Sinai deserts were surveyed for the predominant plant cover and for the culturable bacteria nesting their roots and shoots. Among 43 plant species reported, 13 are perennial (e.g. Fagonia spp., Pancratium spp.) and 30 annuals (e.g. Bromus spp., Erodium spp.). Eleven species possessed rhizo-sheath, e.g. Cyperus capitatus, Panicum turgidum and Trisetaria koelerioides. Microbiological analyses demonstrated: the great diversity and richness of associated culturable bacteria, in particular nitrogen-fixing bacteria (diazotrophs); the majority of bacterial residents were of true and/or putative diazotrophic nature; the bacterial populations followed an increasing density gradient towards the root surfaces; sizeable populations were able to reside inside the root (endorhizosphere) and shoot (endophyllosphere) tissues. Three hundred bacterial isolates were secured from studied spheres. The majority of nitrogen-fixing bacilli isolates belonged to Bacillus megaterium, Bacillus pumilus, Bacillus polymexa, Bacillus macerans, Bacillus circulans and Bacillus licheniformis. The family Enterobacteriaceae represented by Enterobacter agglomerans, Enterobacter sackazakii, Enterobacter cloacae, Serratia adorifera, Serratia liquefaciens and Klebsiella oxytoca. The non-Enterobacteriaceae population was rich in Pantoae spp., Agrobacterium rdiobacter, Pseudomonas vesicularis, Pseudomonas putida, Stenotrophomonas maltophilia, Ochrobactrum anthropi, Sphingomonas paucimobilis and Chrysemonas luteola. Gluconacetobacter diazotrophicus were reported inside root and shoot tissues of a number of tested plants. The dense bacterial populations reported speak well to the very possible significant role played by the endophytic bacterial populations in the survival, in respect of nutrition and health, of existing plants. Such groups of diazotrophs are good candidates, as bio-preparates, to support the growth of future field crops grown in deserts of north Sinai and irrigated by the water of El-Salam canal. [Copyright &y& Elsevier]

We found significant genetic variation in the ability of wheat ( Triticum aestivum) to form rhizosheaths on acid soil and assessed whether differences in aluminium (Al3+) tolerance of root hairs between genotypes was the physiological basis for this genetic variation., A method was developed to rapidly screen rhizosheath size in a range of wheat genotypes. Backcrossed populations were generated from cv Fronteira (large rhizosheath) using cv EGA-Burke (small rhizosheath) as the recurrent parent., A positive correlation existed between rhizosheath size on acid soil and root hair length. In hydroponic experiments, root hairs of the backcrossed lines with large rhizosheaths were more tolerant of Al3+ toxicity than the backcrossed lines with small rhizosheaths., We conclude that greater Al3+ tolerance of root hairs underlies the larger rhizosheath of wheat grown on acid soil. Tolerance of the root hairs to Al3+ was largely independent of the TaALMT1 gene which suggests that different genes encode the Al3+ tolerance of root hairs. The maintenance of longer root hairs in acid soils is important for the efficient uptake of water and nutrients. [ABSTRACT FROM AUTHOR]

Background and Aims Phosphorus commonly limits crop yield and is frequently applied as fertilizer; however, supplies of quality rock phosphate for fertilizer production are diminishing. Plants have evolved many mechanisms to increase their P-fertilizer use efficiency, and an understanding of these traits could result in improved long-term sustainability of agriculture. Here a mutant population is utilized to assess the impact of root hair length on P acquisition and yield under P-deficient conditions alone or when combined with drought. Methods Mutants with various root hair phenotypes were grown in the glasshouse in pots filled with soil representing sufficient and deficient P treatments and, in one experiment, a range of water availability was also imposed. Plants were variously harvested at 7 d, 8 weeks and 14 weeks, and variables including root hair length, rhizosheath weight, biomass, P accumulation and yield were measured. Key Results The results confirmed the robustness of the root hair phenotypes in soils and their relationship to rhizosheath production. The data demonstrated that root hair length is important for shoot P accumulation and biomass, while only the presence of root hairs is critical for yield. Root hair presence was also critical for tolerance to extreme combined P deficit and drought stress, with genotypes with no root hairs suffering extreme growth retardation in comparison with those with root hairs. Conclusions The results suggest that although root hair length is not important for maintaining yield, the presence of root hairs is implicit to sustainable yield of barley under P-deficient conditions and when combined with extreme drought. Root hairs are a trait that should be maintained in future germplasm. [ABSTRACT FROM AUTHOR]

Aims: To illustrate the morphology of sand-binding roots of Haemodoraceae, to conduct a comprehensive survey of the trait, spanning different climates across four continents, and to explore evolutionary hypotheses within a molecular phylogenetic framework. Methods: Sand-binding roots in Haemodoraceae were examined, measured and photographed in the field and on herbarium specimens. Photomicrographs were taken of southwest Australian species. The presence and absence of the sand-binding trait was mapped onto previously published phylogenies and an ancestral state reconstruction was performed. Results: Sand grains were very tightly bound to the root surface by persistent root hairs in Haemodoraceae. The majority of genera and species were found to possess sand-binding roots and only 2 of the 14 genera, Conostylis and Tribonanthes, had sister taxa with and without the trait. The trait was recorded in tropical, sub-tropical and wet temperate species, but mainly in semi-arid species. Sand-binding roots were likely to have been present in the ancestor of the family and both sub-families. Conclusions: The presence of sand-binding roots is the probable ancestral condition for Haemodoraceae, associated with a high degree of phylogenetic conservatism and some secondary loss, notably in Conostylis. Experimental studies are needed to understand the ecological and evolutionary forces at work. [ABSTRACT FROM AUTHOR]

Barley ( Hordeum vulgare L.), genetically modified with the Al3+ resistance gene of wheat ( TaALMT1), was compared with a non-transformed sibling line when grown on an acidic and highly phosphate-fixing ferrosol supplied with a range of phosphorus concentrations. In short-term pot trials (26 days), transgenic barley expressing TaALMT1 (GP-ALMT1) was more efficient than a non-transformed sibling line (GP) at taking up phosphorus on acid soil, but the genotypes did not differ when the soil was limed. Differences in phosphorus uptake efficiency on acid soil could be attributed not only to the differential effects of aluminium toxicity on root growth between the genotypes, but also to differences in phosphorus uptake per unit root length. Although GP-ALMT1 out-performed GP on acid soil, it was still not as efficient at taking up phosphorus as plants grown on limed soil. GP-ALMT1 plants grown in acid soil possessed substantially smaller rhizosheaths than those grown in limed soil, suggesting that root hairs were shorter. This is a probable reason for the lower phosphorus uptake efficiency. When grown to maturity in large pots, GP-ALMT1 plants produced more than twice the grain as GP plants grown on acid soil and 80% of the grain produced by limed controls. Expression of TaALMT1 in barley was not associated with a penalty in either total shoot or grain production in the absence of Al3+, with both genotypes showing equivalent yields in limed soil. These findings demonstrate that an important crop species can be genetically engineered to successfully increase grain production on an acid soil. [ABSTRACT FROM AUTHOR]

The roots of most gramineous plants are surrounded by a variety of microorganisms; however, few studies have focused on the rhizosheath of psammophytes. Therefore, in this study, we used Illumina HiSeq high-throughput sequencing technology to analyse the composition and functional diversity of microbial communities in the rhizosheath of sand-grown Agropyron cristatum (L.) Gaertn. We found that the number of species and functions of microbial communities gradually decreased from the rhizosheath to the bulk soil. Thus, the microbial composition of the rhizosheath was richer and more diverse, and the abundance of bacteria, including Sphingosinicella , Rhizorhabdus , Friedmanniella , Geodermatophilus , Blastococcus , and Oscillatoria , was higher, and the abundance of fungi, such as Mycothermus , was higher. The abundance of CO 2 fixation-related genes ( acsA , Pcc , and cbbL ) in the carbon cycle; NO 3 - , NO 2 - , NH 2 OH, and N 2 transformation genes ( nrtP , nirS , hao , and nifK ) in the nitrogen cycle; soxB / A / C , Sat , and dsrB ) were higher in the rhizosheath than in the bulk soil, as well as genes related to phosphorus uptake in the phosphorus cycle. Our findings showed that the rhizosheath may host the predominant microbial species related to the formation of a rhizosheath.MtlD ) gene and polyketide synthase gene ( pks ) were higher in the rhizosheath than in the bulk soil, as well as genes related to phosphorus uptake in the phosphorus cycle. Our findings showed that the rhizosheath may host the predominant microbial species related to the formation of a rhizosheath.

There is conflicting evidence as to whether rhizosheath soil can be wetter or drier than bulk soil. The gravimetric moisture contents of bulk soil and the rhizosheath of wheat were determined for plants grown in a glasshouse over a range of dry bulk densities (1.0, 1.1 and 1.3 Mg m-3). Plants were also grown in soil at 1. Mg m-2) at 15 °C in a controlled environment. No significant differences in soil moistures over the initial dry bulk density range examined were found. Overall, rhizosheath soil was significantly (P < 0.05) wetter than bulk soil. In the second experiment, under controlled environment conditions, significant differences (P < 0.05) were found only at the lowest depth range examined. The mechanism behind preferential wetting of the rhizosheath is examined with reference to other work on transfer of water to soil from roots, physical changes in the rhizosheath, and root exudates. The most likely explanation of these results is that the presence of mucigel within the rhizosheath increases the water-holding capacity of that soil. [ABSTRACT FROM AUTHOR]

• Rhizosheath influenced the water uptake of drought tolerant chickpea cultivar. • The concentrations of rhizodeposits were low in drought tolerant cultivar. • Rhizosheath pore geometry modified the bacterial abundance at soil-root interface. • The soil-root-microbe interaction affected the water uptake of chickpea cultivars. Improving our understanding of drought tolerance of crops is essential in light of future predicted changes in rainfall, decreased groundwater availability, and increasing temperatures. With a focus on above ground traits, significant improvements in drought tolerance of plants has occurred. With such gains plateauing, we have sought to quantify the belowground functional interactions between plant roots and soil in relation to drought tolerance. Using physical, chemical and biological approaches, we compared drought tolerant and sensitive model plants to demonstrate that a tolerant plant alters both the surrounding pore geometry and the relative abundance of bacteria and upregulates the development of a slow wetting rhizosheath, which increases water uptake under drought conditions. We propose that such rhizosheath traits can be targeted to modify the biophysical properties of the rhizosheath to access water in drought conditions. [ABSTRACT FROM AUTHOR]

The rhizosheath is an important drought-adaptive trait in roots of many angiosperms and has been regarded as a potential trait for future agricultural sustainability. In recent studies, we found that rice roots could form a pronounced rhizosheath under moderate soil drying (MSD) but not under continuous flooding irrigation (CF). The formation of rhizosheaths substantially changes the microbial community structure in endosphere root tissues and the rhizosphere in rice, which may induce a plant immune response. However, the manner by which the formation of rhizosheaths regulates the immune system of roots remains largely unknown. Here, we have analyzed the root transcriptomes of drought-tolerant rice and drought-sensitive rice under both MSD (rhizosheath-root) and CF (root without rhizosheath) conditions. Our results suggest that rhizosheath-associated microbes may trigger plant immune pathways in root under MSD, including the first line of defense component pattern-triggered immunity and the second line of defense component effector-triggered immunity. These data expand our understanding of rhizosheath-associated microbes and plant interactions.

Purpose: Rhizosheath is an adaptive feature for the survival of Stipagrostis pennata in desert systems. Although microorganisms play important ecological roles in promoting the nitrogen cycle of rhizosheath, the diversity and function of nitrogen-fixing microorganism communities have not been fully understood., Materials and Methods: Therefore, the aim of the present study is to explore the nitrogen fixation ability of rhizosheaths and the changes in abundance of nitrogen-fixing microorganisms at different growth periods of S. pennata. We sequenced the nifH gene through sequencing to identify the structure and diversity of nitrogen-fixing microorganisms of S. pennata at different growth periods of rhizosheaths., Results: A total of 1256 operational taxonomic units (OTUs) were identified by nifH sequence and distributed in different growth periods. There were five OTUs distributed in each sample at the same time, and the abundance and diversity of microorganisms in fruit period were much higher than those in other periods. Mainly four phyla were involved, among which Proteobacteria was the most abundant in all groups., Conclusions: In general, the present study investigated the abundance and characteristics of nitrogen-fixing microorganisms of rhizosheaths in different growth periods of S. pennata. It also may elucidate and indicate that the structure of nitrogen-fixing microorganisms of rhizosheaths in different growth periods of S. pennata had changed., (© 2021 The Author(s).)