Root angle is controlled by EGT1 in cereal crops employing an antigravitropic mechanism.

Root angle in crops represents a key trait for efficient capture of soil resources. Root angle is determined by competing gravitropic versus antigravitropic offset (AGO) mechanisms. Here we report a root angle regulatory gene termed ENHANCED GRAVITROPISM1 (EGT1) that encodes a putative AGO component, whose loss-of-function enhances root gravitropism. Mutations in barley and wheat EGT1 genes confer a striking root phenotype, where every root class adopts a steeper growth angle. EGT1 encodes an F-box and Tubby domain-containing protein that is highly conserved across plant species. Haplotype analysis found that natural allelic variation at the barley EGT1 locus impacts root angle. Gravitropic assays indicated that Hvegt1 roots bend more rapidly than wild-type. Transcript profiling revealed Hvegt1 roots deregulate reactive oxygen species (ROS) homeostasis and cell wall-loosening enzymes and cofactors. ROS imaging shows that Hvegt1 root basal meristem and elongation zone tissues have reduced levels. Atomic force microscopy measurements detected elongating Hvegt1 root cortical cell walls are significantly less stiff than wild-type. In situ analysis identified HvEGT1 is expressed in elongating cortical and stele tissues, which are distinct from known root gravitropic perception and response tissues in the columella and epidermis, respectively. We propose that EGT1 controls root angle by regulating cell wall stiffness in elongating root cortical tissue, counteracting the gravitropic machinery's known ability to bend the root via its outermost tissues. We conclude that root angle is controlled by EGT1 in cereal crops employing an antigravitropic mechanism.

Effects of water stress on apoplastic barrier formation in soil grown roots differ from hydroponically grown roots: Histochemical, biochemical and molecular evidence.

In root research, hydroponic plant cultivation is commonly used and soil experiments are rare. We investigated the response of 12-day-old barley roots, cultivated in soil-filled rhizotrons, to different soil water potentials (SWP) comparing a modern cultivar (cv. Scarlett) with a wild accession ICB181243 from Pakistan. Water potentials were quantified in soils with different relative water contents. Root anatomy was studied using histochemistry and microscopy. Suberin and lignin amounts were quantified by analytical chemistry. Transcriptomic changes were observed by RNA-sequencing. Compared with control with decreasing SWP, total root length decreased, the onset of endodermal suberization occurred much closer towards the root tips, amounts of suberin and lignin increased, and corresponding biosynthesis genes were upregulated in response to decreasing SWP. We conclude that decreasing water potentials enhanced root suberization and lignification, like osmotic stress experiments in hydroponic cultivation. However, in soil endodermal cell suberization was initiated very close towards the root tip, and root length as well as suberin amounts were about twofold higher compared with hydroponic cultivation.

ENHANCED GRAVITROPISM 2 encodes a STERILE ALPHA MOTIF-containing protein that controls root growth angle in barley and wheat.

The root growth angle defines how roots grow toward the gravity vector and is among the most important determinants of root system architecture. It controls water uptake capacity, nutrient use efficiency, stress resilience, and, as a consequence, yield of crop plants. We demonstrated that the egt2 (enhanced gravitropism 2) mutant of barley exhibits steeper root growth of seminal and lateral roots and an auxin-independent higher responsiveness to gravity compared to wild-type plants. We cloned the EGT2 gene by a combination of bulked-segregant analysis and whole genome sequencing. Subsequent validation experiments by an independent CRISPR/Cas9 mutant allele demonstrated that egt2 encodes a STERILE ALPHA MOTIF domain-containing protein. In situ hybridization experiments illustrated that EGT2 is expressed from the root cap to the elongation zone. We demonstrated the evolutionary conserved role of EGT2 in root growth angle control between barley and wheat by knocking out the EGT2 orthologs in the A and B genomes of tetraploid durum wheat. By combining laser capture microdissection with RNA sequencing, we observed that seven expansin genes were transcriptionally down-regulated in the elongation zone. This is consistent with a role of EGT2 in this region of the root where the effect of gravity sensing is executed by differential cell elongation. Our findings suggest that EGT2 is an evolutionary conserved regulator of root growth angle in barley and wheat that could be a valuable target for root-based crop improvement strategies in cereals.

The root system architecture of wheat establishing in soil is associated with varying elongation rates of seminal roots: quantification using 4D magnetic resonance imaging.

Seedling establishment is the first stage of crop productivity, and root phenotypes at seed emergence are critical to a successful start of shoot growth as well as for water and nutrient uptake. In this study, we investigate seedling establishment in winter wheat utilizing a newly developed workflow based on magnetic resonance imaging (MRI). Using the eight parents of the MAGIC (multi-parent advanced generation inter-cross) population we analysed the 4D root architecture of 288 individual seedlings grown in natural soils with plant neighbors over 3 d of development. Time of root and shoot emergence, total length, angle, and depth of the axile roots varied significantly among these genotypes. The temporal data resolved rates of elongation of primary roots and first and second seminal root pairs. Genotypes with slowly elongating primary roots had rapidly elongating first and second seminal root pairs and vice versa, resulting in variation in root system architecture mediated not only by root angle but also by initiation and relative elongation of axile roots. We demonstrated that our novel MRI workflow with a unique planting design and automated measurements allowed medium throughput phenotyping of wheat roots in 4D and could give new insights into regulation of root system architecture.

Root architecture simulation improves the inference from seedling root phenotyping towards mature root systems.

Root phenotyping provides trait information for plant breeding. A shortcoming of high-throughput root phenotyping is the limitation to seedling plants and failure to make inferences on mature root systems. We suggest root system architecture (RSA) models to predict mature root traits and overcome the inference problem. Sixteen pea genotypes were phenotyped in (i) seedling (Petri dishes) and (ii) mature (sand-filled columns) root phenotyping platforms. The RSA model RootBox was parameterized with seedling traits to simulate the fully developed root systems. Measured and modelled root length, first-order lateral number, and root distribution were compared to determine key traits for model-based prediction. No direct relationship in root traits (tap, lateral length, interbranch distance) was evident between phenotyping systems. RootBox significantly improved the inference over phenotyping platforms. Seedling plant tap and lateral root elongation rates and interbranch distance were sufficient model parameters to predict genotype ranking in total root length with an RSpearman of 0.83. Parameterization including uneven lateral spacing via a scaling function substantially improved the prediction of architectures underlying the differently sized root systems. We conclude that RSA models can solve the inference problem of seedling root phenotyping. RSA models should be included in the phenotyping pipeline to provide reliable information on mature root systems to breeding research.

Screening for drought tolerance of maize hybrids by multi-scale analysis of root and shoot traits at the seedling stage.

We studied the drought response of eight commercial hybrid maize lines with contrasting drought sensitivity together with the reference inbred line B73 using a non-invasive platform for root and shoot phenotyping and a kinematics approach to quantify cell level responses in the leaf. Drought treatments strongly reduced leaf growth parameters including projected leaf area, elongation rate, final length and width of the fourth and fifth leaf. Physiological measurements including water use efficiency, chlorophyll fluorescence and photosynthesis were also significantly affected. By performing a kinematic analysis, we show that leaf growth reduction in response to drought is mainly due to a decrease in cell division rate, whereas a marked reduction in cell expansion rate is compensated by increased duration of cell expansion. Detailed analysis of root growth in rhizotrons under drought conditions revealed a strong reduction in total root length as well as rooting depth and width. This was reflected by corresponding decreases in fresh and dry weight of the root system. We show that phenotypic differences between lines differing in geographic origin (African vs. European) and in drought tolerance under field conditions can already be identified at the seedling stage by measurements of total root length and shoot dry weight of the plants. Moreover, we propose a list of candidate traits that could potentially serve as traits for future screening strategies.

Plant root tortuosity: an indicator of root path formation in soil with different composition and density.

Background and Aims Root soil penetration and path optimization are fundamental for root development in soil. We describe the influence of soil strength on root elongation rate and diameter, response to gravity, and root-structure tortuosity, estimated by average curvature of primary maize roots. Methods Soils with different densities (1·5, 1·6, 1·7 g cm-3), particle sizes (sandy loam; coarse sand mixed with sandy loam) and layering (monolayer, bilayer) were used. In total, five treatments were performed: Mix_low with mixed sand low density (three pots, 12 plants), Mix_medium - mixed sand medium density (three pots, 12 plants), Mix_high - mixed sand high density (three pots, ten plants), Loam_low sandy loam soil low density (four pots, 16 plants), and Bilayer with top layer of sandy loam and bottom layer mixed sand both of low density (four pots, 16 plants). We used non-invasive three-dimensional magnetic resonance imaging to quantify effects of these treatments. Key Results Roots grew more slowly [root growth rate (mm h-1); decreased 50 %] with increased diameters [root diameter (mm); increased 15 %] in denser soils (1·7 vs. 1·5 g cm-3). Root response to gravity decreased 23 % with increased soil compaction, and tortuosity increased 10 % in mixed sand. Response to gravity increased 39 % and tortuosity decreased 3 % in sandy loam. After crossing a bilayered-soil interface, roots grew more slowly, similar to roots grown in soil with a bulk density of 1·64 g cm-3, whereas the actual experimental density was 1·48±0·02 g cm-3. Elongation rate and tortuosity were higher in Mix_low than in Loam_low. Conclusions The present study increases our existing knowledge of the influence of physical soil properties on root growth and presents new assays for studying root growth dynamics in non-transparent media. We found that root tortuosity is indicative of root path selection, because it could result from both mechanical deflection and active root growth in response to touch stimulation and mechanical impedance.

Cooperative action of the paralogous maize lateral organ boundaries (LOB) domain proteins RTCS and RTCL in shoot-borne root formation.

The paralogous maize (Zea mays) LBD (Lateral Organ Boundaries Domain) genes rtcs (rootless concerning crown and seminal roots) and rtcl (rtcs-like) emerged from an ancient whole-genome duplication. RTCS is a key regulator of crown root initiation. The diversity of expression, molecular interaction and phenotype of rtcs and rtcl were investigated. The rtcs and rtcl genes display highly correlated spatio-temporal expression patterns in roots, despite the significantly higher expression of rtcs. Both RTCS and RTCL proteins bind to LBD downstream promoters and act as transcription factors. In line with its auxin inducibility and binding to auxin response elements of rtcs and rtcl promoters, ARF34 (AUXIN RESPONSE FACTOR 34) acts as transcriptional activator. Yeast two-hybrid screening combined with bimolecular fluorescence complementation (BiFC) experiments revealed conserved and unique interaction partners of RTCS and RTCL. The rtcl mutation leads to defective shoot-borne root elongation early in development. Cooperative action of RTCS and RTCL during shoot-borne root formation was demonstrated by rtcs-dependent repression of rtcl transcription in coleoptilar nodes. Although RTCS is instrumental in shoot-borne root initiation, RTCL controls shoot-borne root elongation early in development. Their conserved role in auxin signaling, but diverse function in shoot-borne root formation, is underscored by their conserved and unique interaction partners.

Simultaneous effects of leaf irradiance and soil moisture on growth and root system architecture of novel wheat genotypes: implications for phenotyping.

Plants in the field are exposed to varying light and moisture. Agronomic improvement requires knowledge of whole-plant phenotypes expressed in response to simultaneous variation in these essential resources. Most phenotypes, however, have been described from experiments where resources are varied singularly. To test the importance of varying shoot and root resources for phenotyping studies, sister pre-breeding lines of wheat were phenotyped in response to independent or simultaneous exposure to two light levels and soil moisture profiles. The distribution and architecture of the root systems depended strongly on the moisture of the deeper soil layer. For one genotype, roots, specifically lateral roots, were stimulated to grow into moist soil when the upper zone was well-watered and were inhibited by drier deep zones. In contrast, the other genotype showed much less plasticity and responsiveness to upper moist soil, but maintained deeper penetration of roots into the dry layer. The sum of shoot and root responses was greater when treated simultaneously to low light and low soil water, compared to each treatment alone, suggesting the value of whole plant phenotyping in response to multiple conditions for agronomic improvement. The results suggest that canopy management for increased irradiation of leaves would encourage root growth into deeper drier soil, and that genetic variation within closely related breeding lines may exist to favour surface root growth in response to irrigation or in-season rainfall.

Impact of domestication on the phenotypic architecture of durum wheat under contrasting nitrogen fertilization.

The process of domestication has led to dramatic morphological and physiological changes in crop species due to adaptation to cultivation and to the needs of farmers. To investigate the phenotypic architecture of shoot- and root-related traits and quantify the impact of primary and secondary domestication, we examined a collection of 36 wheat genotypes under optimal and nitrogen-starvation conditions. These represented three taxa that correspond to key steps in the recent evolution of tetraploid wheat (i.e. wild emmer, emmer, and durum wheat). Overall, nitrogen starvation reduced the shoot growth of all genotypes, while it induced the opposite effect on root traits, quantified using the automated phenotyping platform GROWSCREEN-Rhizo. We observed an overall increase in all of the shoot and root growth traits from wild emmer to durum wheat, while emmer was generally very similar to wild emmer but intermediate between these two subspecies. While the differences in phenotypic diversity due to the effects of primary domestication were not significant, the secondary domestication transition from emmer to durum wheat was marked by a large and significant decrease in the coefficient of additive genetic variation. In particular, this reduction was very strong under the optimal condition and less intense under nitrogen starvation. Moreover, although under the optimal condition both root and shoot traits showed significantly reduced diversity due to secondary domestication, under nitrogen starvation the reduced diversity was significant only for shoot traits. Overall, a considerable amount of phenotypic variation was observed in wild emmer and emmer, which could be exploited for the development of pre-breeding strategies.

Phenotype of Arabidopsis thaliana semi-dwarfs with deep roots and high growth rates under water-limiting conditions is independent of the GA5 loss-of-function alleles.

BACKGROUND AND AIMS: The occurrence of Arabidopsis thaliana semi-dwarf accessions carrying inactive alleles at the gibberellin (GA) biosynthesis GA5 locus has raised the question whether there are pleiotropic effects on other traits at the root level, such as rooting depth. In addition, it is unknown whether semi-dwarfism in arabidopsis confers a growth advantage under water-limiting conditions compared with wild-type plants. The aim of this research was therefore to investigate whether semi-dwarfism has a pleiotropic effect in the root system and also whether semi-dwarfs might be more tolerant of water-limiting conditions. METHODS: The root systems of different arabidopsis semi-dwarfs and GA biosynthesis mutants were phenotyped in vitro using the GROWSCREEN-ROOT image-based software. Semi-dwarfs were phenotyped together with tall, near-related accessions. In addition, root phenotypes were investigated in soil-filled rhizotrons. Rosette growth trajectories were analysed with the GROWSCREEN-FLUORO setup based on non-invasive imaging. KEY RESULTS: Mutations in the early steps of the GA biosynthesis pathway led to a reduction in shoot as well as root size. Depending on the genetic background, mutations at the GA5 locus yielded phenotypes characterized by decreased root length in comparison with related wild-type ones. The semi-dwarf accession Pak-3 showed the deepest root system both in vitro and in soil cultivation experiments; this comparatively deep root system, however, was independent of the ga5 loss-of-function allele, as shown by co-segregation analysis. When the accessions were grown under water-limiting conditions, semi-dwarf accessions with high growth rates were identified. CONCLUSIONS: The observed diversity in root system growth and architecture occurs independently of semi-dwarf phenotypes, and is probably linked to a genetic background effect. The results show that there are no clear advantages of semi-dwarfism at low water availability in arabidopsis.

Precrop-treated soil influences wheat (Triticum aestivum L.) root system architecture and its response to drought.

Aims: Root system architecture (RSA) plays an important role in the plant's ability to sustain yield under abiotic stresses such as drought. Preceding crops (precrops) can affect the yield of the proceeding crop, partially by affecting the RSA. This experiment aims to explore the interactions between precrop identity, crop genotype and drought at early growth stages. Methods: Rhizotrons, sized 60 × 80 × 3.5 cm, were used to assess the early root growth of two winter wheat (Triticum aestivum L.) genotypes, using precrop-treated soil around the seedlings and differing water regimes. The rhizotrons were automatically imaged 3 times a week to track root development. Results: Precrop-treated soil affected the RSA and changes caused by the reduced water treatment (RWT) were different depending on the precrop. Largest of these was the 36% reduction in root depth after wheat, but 44% after OSR. This indicates that effects caused by the precrop can be simulated, at least partially, by transferring precrop-treated soils to controlled environments. The genotypes had differential RSA and reacted differently to the RWT, with Julius maintaining an 8.8-13.1% deeper root system compared to Brons in the RWT. In addition, the combined environmental treatment affected the genotypes differently. Conclusion: Our results could help explain discrepancies found from using precrops to enhance yield as they indicate differences in the preceding crop effect when experiencing drought stress. Further, these differences are affected by genotypic interactions, which can be used to select and adapt crop genotypes for specific crop rotations, depending on the year. Additionally, we have shown a viable method of stimulating a partial precrop effect at the seedling stage in a controlled greenhouse setting using field soil around the germinated seed.

Effects of altered α- and ß-branch carotenoid biosynthesis on photoprotection and whole-plant acclimation of Arabidopsis to photo-oxidative stress.

Functions of α- and ß-branch carotenoids in whole-plant acclimation to photo-oxidative stress were studied in Arabidopsis thaliana wild-type (wt) and carotenoid mutants, lutein deficient (lut2, lut5), non-photochemical quenching1 (npq1) and suppressor of zeaxanthin-less1 (szl1) npq1 double mutant. Photo-oxidative stress was applied by exposing plants to sunflecks. The sunflecks caused reduction of chlorophyll content in all plants, but more severely in those having high α- to ß-branch carotenoid composition (α/ß-ratio) (lut5, szl1npq1). While this did not alter carotenoid composition in wt or lut2, which accumulates only ß-branch carotenoids, increased xanthophyll levels were found in the mutants with high α/ß-ratios (lut5, szl1npq1) or without xanthophyll-cycle operation (npq1, szl1npq1). The PsbS protein content increased in all sunfleck plants but lut2. These changes were accompanied by no change (npq1, szl1npq1) or enhanced capacity (wt, lut5) of NPQ. Leaf mass per area increased in lut2, but decreased in wt and lut5 that showed increased NPQ. The sunflecks decelerated primary root growth in wt and npq1 having normal α/ß-ratios, but suppressed lateral root formation in lut5 and szl1npq1 having high α/ß-ratios. The results highlight the importance of proper regulation of the α- and ß-branch carotenoid pathways for whole-plant acclimation, not only leaf photoprotection, under photo-oxidative stress.

Root-root interactions: extending our perspective to be more inclusive of the range of theories in ecology and agriculture using in-vivo analyses.

BACKGROUND: There is a large body of literature on competitive interactions among plants, but many studies have only focused on above-ground interactions and little is known about root-root dynamics between interacting plants. The perspective on possible mechanisms that explain the outcome of root-root interactions has recently been extended to include non-resource-driven mechanisms (as well as resource-driven mechanisms) of root competition and positive interactions such as facilitation. These approaches have often suffered from being static, partly due to the lack of appropriate methodologies for in-situ non-destructive root characterization. SCOPE: Recent studies show that interactive effects of plant neighbourhood interactions follow non-linear and non-additive paths that are hard to explain. Common outcomes such as accumulation of roots mainly in the topsoil cannot be explained solely by competition theory but require a more inclusive theoretical, as well as an improved methodological framework. This will include the question of whether we can apply the same conceptual framework to crop versus natural species. CONCLUSIONS: The development of non-invasive methods to dynamically study root-root interactions in vivo will provide the necessary tools to study a more inclusive conceptual framework for root-root interactions. By following the dynamics of root-root interactions through time in a whole range of scenarios and systems, using a wide variety of non-invasive methods, (such as fluorescent protein which now allows us to separately identify the roots of several individuals within soil), we will be much better equipped to answer some of the key questions in root physiology, ecology and agronomy.

Pitfalls and potential of high-throughput plant phenotyping platforms.

Automated high-throughput plant phenotyping (HTPP) enables non-invasive, fast and standardized evaluations of a large number of plants for size, development, and certain physiological variables. Many research groups recognize the potential of HTPP and have made significant investments in HTPP infrastructure, or are considering doing so. To make optimal use of limited resources, it is important to plan and use these facilities prudently and to interpret the results carefully. Here we present a number of points that users should consider before purchasing, building or utilizing such equipment. They relate to (1) the financial and time investment for acquisition, operation, and maintenance, (2) the constraints associated with such machines in terms of flexibility and growth conditions, (3) the pros and cons of frequent non-destructive measurements, (4) the level of information provided by proxy traits, and (5) the utilization of calibration curves. Using data from an Arabidopsis experiment, we demonstrate how diurnal changes in leaf angle can impact plant size estimates from top-view cameras, causing deviations of more than 20% over the day. Growth analysis data from another rosette species showed that there was a curvilinear relationship between total and projected leaf area. Neglecting this curvilinearity resulted in linear calibration curves that, although having a high r2 (> 0.92), also exhibited large relative errors. Another important consideration we discussed is the frequency at which calibration curves need to be generated and whether different treatments, seasons, or genotypes require distinct calibration curves. In conclusion, HTPP systems have become a valuable addition to the toolbox of plant biologists, provided that these systems are tailored to the research questions of interest, and users are aware of both the possible pitfalls and potential involved.

Unraveling Genomic Regions Controlling Root Traits as a Function of Nitrogen Availability in the MAGIC Wheat Population WM-800.

An ever-growing world population demands to be fed in the future and environmental protection and climate change need to be taken into account. An important factor here is nitrogen uptake efficiency (NUpE), which is influenced by the root system (the interface between plant and soil). To understand the natural variation of root system architecture (RSA) as a function of nitrogen (N) availability, a subset of the multiparent advanced generation intercross (MAGIC) winter wheat population WM-800 was phenotyped under two contrasting N treatments in a high-throughput phenotyping system at the seedling stage. Fourteen root and shoot traits were measured. Subsequently, these traits were genetically analyzed using 13,060 polymorphic haplotypes and SNPs in a genome-wide association study (GWAS). In total, 64 quantitative trait loci (QTL) were detected; 60 of them were N treatment specific. Candidate genes for the detected QTL included NRT1.1 and genes involved in stress signaling under N-, whereas candidate genes under N+ were more associated with general growth, such as mei2 and TaWOX11b. This finding may indicate (i) a disparity of the genetic control of root development under low and high N supply and, furthermore, (ii) the need for an N specific selection of genes and genotypes in breeding new wheat cultivars with improved NUpE.

Co-fertilization of Sulfur and Struvite-Phosphorus in a Slow-Release Fertilizer Improves Soybean Cultivation.

In face of the alarming world population growth predictions and its threat to food security, the development of sustainable fertilizer alternatives is urgent. Moreover, fertilizer performance should be assessed not only in terms of yield but also in root system development, as it impacts soil fertility and crop productivity. Fertilizers containing a polysulfide matrix (PS) with dispersed struvite (St) were studied for S and P nutrition due to their controlled-release behavior. Soybean cultivation in a closed system with St/PS composites provided superior biomass compared to a reference of triple superphosphate (TSP) with ammonium sulfate (AS), with up to 3 and 10 times higher mass of shoots and roots, respectively. Root system architectural changes may explain these results, with a higher proliferation of second order lateral roots in response to struvite ongoing P delivery. The total root length was between 1,942 and 4,291 cm for plants under St/PS composites and only 982 cm with TSP/AS. While phosphorus uptake efficiency was similar in all fertilized treatments (11-14%), St/PS achieved a 22% sulfur uptake efficiency against only 8% from TSP/AS. Overall, the composites showed great potential as efficient slow-release fertilizers for enhanced soybean productivity.

Plant Growth Promotion and Heat Stress Amelioration in Arabidopsis Inoculated with Paraburkholderia phytofirmans PsJN Rhizobacteria Quantified with the GrowScreen-Agar II Phenotyping Platform.

High temperatures inhibit plant growth. A proposed strategy for improving plant productivity under elevated temperatures is the use of plant growth-promoting rhizobacteria (PGPR). While the effects of PGPR on plant shoots have been extensively explored, roots-particularly their spatial and temporal dynamics-have been hard to study, due to their below-ground nature. Here, we characterized the time- and tissue-specific morphological changes in bacterized plants using a novel non-invasive high-resolution plant phenotyping and imaging platform-GrowScreen-Agar II. The platform uses custom-made agar plates, which allow air exchange to occur with the agar medium and enable the shoot to grow outside the compartment. The platform provides light protection to the roots, the exposure of it to the shoots, and the non-invasive phenotyping of both organs. Arabidopsis thaliana, co-cultivated with Paraburkholderia phytofirmans PsJN at elevated and ambient temperatures, showed increased lengths, growth rates, and numbers of roots. However, the magnitude and direction of the growth promotion varied depending on root type, timing, and temperature. The root length and distribution per depth and according to time was also influenced by bacterization and the temperature. The shoot biomass increased at the later stages under ambient temperature in the bacterized plants. The study offers insights into the timing of the tissue-specific, PsJN-induced morphological changes and should facilitate future molecular and biochemical studies on plant-microbe-environment interactions.

Recovery dynamics of growth, photosynthesis and carbohydrate accumulation after de-submergence: a comparison between two wetland plants showing escape and quiescence strategies.

BACKGROUND AND AIMS: The capacity for fast-growth recovery after de-submergence is important for establishment of riparian species in a water-level-fluctuation zone. Recovery patterns of two wetland plants, Alternanthera philoxeroides and Hemarthria altissima, showing 'escape' and 'quiescence' responses, respectively, during submergence were investigated. METHODS: Leaf and root growth and photosynthesis were monitored continuously during 10 d of recovery following 20 d of complete submergence. Above- and below-ground dry weights, as well as carbohydrate concentrations, were measured several times during the experiment. KEY RESULTS: Both species remobilized stored carbohydrate during submergence. Although enhanced internode elongation depleted the carbohydrate storage in A. philoxeroides during submergence, this species resumed leaf growth 3 d after de-submergence concomitant with restoration of the maximal photosynthetic capacity. In contrast, some sucrose was conserved in shoots of H. altissima during submergence, which promoted rapid re-growth of leaves 2 d after de-submergence and earlier than the full recovery of photosynthesis. The recovery of root growth was delayed by 1-2 d compared with leaves in both species. CONCLUSIONS: Submergence tolerance of the escape and quiescence strategies entails not only the corresponding regulation of growth, carbohydrate catabolism and energy metabolism during submergence but also co-ordinated recovery of photosynthesis, growth and carbohydrate partitioning following de-submergence.

Deep soil exploration vs. topsoil exploitation: distinctive rooting strategies between wheat landraces and wild relatives.

AIMS: Diversity of root systems among genetic resources can contribute to optimize water and nutrient uptake. Topsoil exploitation vs. deep soil exploration represent two contrasting ideotypes in relation to resource use. Our study reveals how rooting patterns changed between wheat wild progenitors and landraces in regard to these ideotypes. METHODS: Root (partitioning, morphology, distribution, elongation, anatomy) and shoot traits (dry-matter, leaf area, assimilation) of durum landraces, wild emmer and wild einkorn from Iran, Syria, Turkey and Lebanon were phenotyped using the GrowScreen-Rhizo platform. Distinctive rooting patterns were identified via principal component analysis and relations with collection site characteristics analyzed. RESULTS: Shoot trait differentiation was strongly driven by seed weight, leading to superior early vigor of landraces. Wild progenitors formed superficial root systems with a higher contribution of lateral and early-emerging nodal axes to total root length. Durum landraces had a root system dominated by seminal axes allocated evenly over depth. Xylem anatomy was the trait most affected by the environmental influence of the collection site. CONCLUSIONS: The durum landrace root system approximated a deep soil exploration ideotype which would optimize subsoil water uptake, while monococcum-type wild einkorn was most similar to a topsoil exploiting strategy with potential competitive advantages for subsistence in natural vegetation. SUPPLEMENTARY INFORMATION: The online version contains supplementary material available at 10.1007/s11104-020-04794-9.