Anatomical and physiological responses of roots and rhizomes in Oryza longistaminata to soil water gradients.

BACKGROUND AND AIMS: Roots and rhizomes are critical for the adaptation of clonal plants to soil water gradients. Oryza longistaminata, a rhizomatous wild rice, is of particular interest for perennial rice breeding due to its resilience under abiotic stress conditions. While root responses to soil flooding are well-studied, rhizome responses to water gradients remain underexplored. We hypothesize that physiological integration of Oryza longistaminata mitigates heterogeneous water deficit stress through interconnected rhizomes, and both roots and rhizomes respond to contrasting water conditions. METHODS: We investigated the physiological integration between mother plants and ramets, measuring key photosynthetic parameters (photosynthetic and transpiration rate, and stomatal conductance) using an Infrared Gas Analyzer. Moreover, root and rhizome responses to three water regimes (flooding, well-watered, and water deficit) were examined by measuring radial water loss and apparent permeance to O2, along with histochemical and anatomical characterization. KEY RESULTS: Our experiment highlights the role of physiological integration via interconnected rhizomes in mitigating water deficit stress. Severing rhizome connections from mother plants or ramets exposed to water deficit conditions led to significant decreases in key photosynthetic parameters, underscoring the importance of rhizome connections in bidirectional stress mitigation. Additionally, O. longistaminata rhizomes exhibited constitutive suberized and lignified apoplastic barriers, while such barriers were induced in roots under water stress. Anatomically, both rhizomes and roots respond similarly to water gradients, showing thinner diameters under water deficit conditions and larger diameters under flooding conditions. CONCLUSION: Our findings indicate that physiological integration through interconnected rhizomes helps alleviate water deficit stress when either the mother plant or the ramet is experiencing water deficit, while the counterpart is in control conditions. Moreover, O. longistaminata can adapt to various soil water regimes by regulating anatomical and physiological traits of roots and rhizomes.

Coordination of pinna, petiole, and root anatomical traits in 24 tropical-subtropical fern species.

Ferns are primitive vascular plants with diverse morphologies and structures. Plant anatomical traits and their linkages can reflect adaptation to the environment; however, these remain are still poorly understood in ferns. The main objective of this study was to explore whether there was structural coordination among and within organs in fern species. We measured 16 hydraulically related anatomical traits of pinnae, petioles, and roots of 24 representative fern species from the tropical and subtropical forest understory and analyzed trait correlation networks. In addition, we examined phylogenetic signals for the anatomical traits and analyzed co-evolutionary relationships. These results indicated that stomatal density and all petiole anatomical traits exhibited significant phylogenetic signals. Evolutionary correlations were observed between the tracheid diameter and wall thickness of the petiole and between the water transport capacity of the petiole and stomatal density. Conversely, anatomical traits of roots (e.g., root diameter) showed no phylogenetic signals and were not significantly correlated with those of the pinnae and petioles, indicating a lack of structural coordination between the below- and above-ground organs. Unlike angiosperms, vein density is unrelated to stomatal density or pinna thickness in ferns. As root diameter decreased, the cortex-to-stele diameter ratio decreased significantly (enhanced water absorption) in angiosperms but remained unchanged in ferns. These differences lead to different responses of ferns to climate change and improve our knowledge of the water adaptation strategies of ferns.

From cells to stems: the effects of primary vascular construction on drought-induced embolism in fern rhizomes.

While a considerable amount of data exists on the link between xylem construction and hydraulic function, few studies have focused on resistance to drought-induced embolism of primary vasculature in herbaceous plants. Ferns rely entirely on primary xylem and display a remarkable diversity of vascular construction in their rhizomes, making them an ideal group in which to examine hydraulic structure-function relationships. New optical methods allowed us to measure vulnerability to embolism in rhizomes, which are notoriously difficult to work with. We investigated five fern species based on their diverse xylem traits at the cellular, histological, and architectural levels. To link below- and above-ground hydraulics, we then measured leaf-stem vulnerability segmentation. Overall, rhizome vulnerability to embolism was correlated most strongly with cellular but not histological or architectural traits. Interestingly, at P6-12 , species with increased architectural dissection were actually more vulnerable to embolism, suggesting different hydraulic dynamics at low compared to high percent embolism. Importantly, leaves fully embolize before stems reach P88 , suggesting strong vulnerability segmentation. This is the first study to explore the functional implications of primary vascular construction in fern rhizomes and leaf-stem vulnerability segmentation. Strong segmentation suggests that leaves protect perennial rhizomes against severe drought stress and hydraulically induced mortality.

Key root traits of Poaceae for adaptation to soil water gradients.

Drought and flooding are contrasting abiotic stressors for plants. Evidence is accumulating for root anatomical traits being essential for the adaptation to drought or flooding. However, an integrated approach to comprehensively understand root anatomical traits has not yet been established. Here we analysed the root anatomical traits of 18 wild Poaceae species differing in adaptation to a range of soil water content. Regression model analyses revealed the optimal anatomical traits that were required by the plants to adapt to low or high soil water content. While the area and number of each root tissue (e.g. stele, cortex, xylem or aerenchyma) were not strongly correlated to the soil water content, the ratio of the root tissue areas (cortex to stele ratio (CSR), xylem to stele ratio (XSR) and aerenchyma to cortex ratio (ACR)) could fully explain the adaptations of the wild Poaceae species to the soil water gradients. Our results demonstrate that the optimal anatomical traits for the adaptations to soil water content can be determined by three indices (i.e. CSR, XSR and ACR), and thus we propose that these root anatomical indices can be used to improve the tolerance of crops to drought and flooding stresses.

Anatomical patterns of condensed tannin in fine roots of tree species from a cool-temperate forest.

BACKGROUND AND AIMS: Condensed tannin (CT) is an important compound in plant biological structural defence and for tolerance of herbivory and environmental stress. However, little is known of the role and location of CT within the fine roots of woody plants. To understand the role of CT in fine roots across diverse species of woody dicot, we evaluated the localization of CT that accumulated in root tissue, and examined its relationships with the stele and cortex tissue in cross-sections of roots in 20 tree species forming different microbial symbiotic groups (ectomycorrhiza and arbuscular mycorrhiza). METHODS: In a cool-temperate forest in Japan, cross-sections of sampled roots in different branching order classes, namely, first order, second to third order, fourth order, and higher than fourth order (higher order), were measured in terms of the length-based ratios of stele diameter and cortex thickness to root diameter. All root samples were then stained with ρ-dimethylaminocinnamaldehyde solution and we determined the ratio of localized CT accumulation area to the root cross-section area (CT ratio). KEY RESULTS: Stele ratio tended to increase with increasing root order, whereas cortex ratio either remained unchanged or decreased with increasing order in all species. The CT ratio was significantly positively correlated to the stele ratio and negatively correlated to the cortex ratio in second- to fourth-order roots across species during the shift from primary to secondary root growth. Ectomycorrhiza-associated species mostly had a higher stele ratio and lower cortex ratio than arbuscular mycorrhiza-associated species across root orders. Compared with arbuscular mycorrhiza species, there was greater accumulation of CT in response to changes in the root order of ectomycorrhiza species. CONCLUSIONS: Different development patterns of the stele, cortex and CT accumulation along the transition from root tip to secondary roots could be distinguished between different mycorrhizal associations. The CT in tissues in different mycorrhizal associations could help with root protection in specific branching orders during shifts in stele and cortex development before and during cork layer formation.

A new phylogeny of the cladoxylopsid plexus: contribution of an early cladoxylopsid from the Lower Devonian (Emsian) of Quebec.

PREMISE: Cladoxylopsids formed Earth's earliest forests and gave rise to the ancestors of sphenopsids and ferns. Lower Devonian (Emsian) strata of the Battery Point Formation (Quebec, Canada) contain new anatomically preserved cladoxylopsids, one of which is described in this article. To assess the phylogenetic position of this fossil and address questions of cladoxylopsid phylogeny, we conducted a comprehensive phylogenetic study. METHODS: Permineralized axes were studied in serial sections using the cellulose acetate peel technique. We evaluated phylogenetic relationships among cladoxylopsids using a data set of 36 new morphological characters and 31 species, in parsimony-constrained analyses. RESULTS: We describe Adelocladoxis praecox gen. et sp. nov., a cladoxylopsid with small actinostelic axes bearing dichotomously branched, helically arranged ultimate appendages and fusiform sporangia. Adelocladoxis provides the oldest evidence of cladoxylopsid anatomy, including ultimate appendages and sporangia. In agreement with non-phylogenetic classification schemes, our phylogenetic analysis resolves a basal grade of iridopterids and a clade of cladoxylopsids s.s., which includes a pseudosporochnalean cladoxylopsid clade, a cladoxylalean cladoxylopsid clade, and Adelocladoxis. CONCLUSIONS: Our phylogenetic analysis illuminates aspects of tempo and mode of evolution in the cladoxylopsid plexus. Originating prior to the Emsian, cladoxylopsids reached global distribution by the Frasnian. Iridopterids and cladoxylopsids s.s. radiated in the Emsian-Eifelian. The sequence of character change recovered by our phylogeny supports a transition from actinostelic protosteles to dissected steles, associated with an increase in xylem rib number and medullation generating a central parenchymatous area.

Climate-smart crops: key root anatomical traits that confer flooding tolerance.

Plants require water, but a deficit or excess of water can negatively impact their growth and functioning. Soil flooding, in which root-zone is filled with excess water, restricts oxygen diffusion into the soil. Global climate change is increasing the risk of crop yield loss caused by flooding, and the development of flooding tolerant crops is urgently needed. Root anatomical traits are essential for plants to adapt to drought and flooding, as they determine the balance between the rates of water and oxygen transport. The stele contains xylem and the cortex contains aerenchyma (gas spaces), which respectively contribute to water uptake from the soil and oxygen supply to the roots; this implies that there is a trade-off between the ratio of cortex and stele sizes with respect to adaptation to drought or flooding. In this review, we analyze recent advances in the understanding of root anatomical traits that confer drought and/or flooding tolerance to plants and illustrate the trade-off between cortex and stele sizes. Moreover, we introduce the progress that has been made in modelling and fully automated analyses of root anatomical traits and discuss how key root anatomical traits can be used to improve crop tolerance to soil flooding.

Comparing Kinetics of Xylem Ion Loading and Its Regulation in Halophytes and Glycophytes.

Although control of xylem ion loading is essential to confer salinity stress tolerance, specific details behind this process remain elusive. In this work, we compared the kinetics of xylem Na+ and K+ loading between two halophytes (Atriplex lentiformis and quinoa) and two glycophyte (pea and beans) species, to understand the mechanistic basis of the above process. Halophyte plants had high initial amounts of Na+ in the leaf, even when grown in the absence of the salt stress. This was matched by 7-fold higher xylem sap Na+ concentration compared with glycophyte plants. Upon salinity exposure, the xylem sap Na+ concentration increased rapidly but transiently in halophytes, while in glycophytes this increase was much delayed. Electrophysiological experiments using the microelectrode ion flux measuring technique showed that glycophyte plants tend to re-absorb Na+ back into the stele, thus reducing xylem Na+ load at the early stages of salinity exposure. The halophyte plants, however, were capable to release Na+ even in the presence of high Na+ concentrations in the xylem. The presence of hydrogen peroxide (H2O2) [mimicking NaCl stress-induced reactive oxygen species (ROS) accumulation in the root] caused a massive Na+ and Ca2+ uptake into the root stele, while triggering a substantial K+ efflux from the cytosol into apoplast in glycophyte but not halophytes species. The peak in H2O2 production was achieved faster in halophytes (30 min vs 4 h) and was attributed to the increased transcript levels of RbohE. Pharmacological data suggested that non-selective cation channels are unlikely to play a major role in ROS-mediated xylem Na+ loading.

Root O2 consumption, CO2 production and tissue concentration profiles in chickpea, as influenced by environmental hypoxia.

Roots in flooded soils experience hypoxia, with the least O2 in the vascular cylinder. Gradients in CO2 across roots had not previously been measured. The respiratory quotient (RQ; CO2 produced : O2 consumed) is expected to increase as O2 availability declines. A new CO2 microsensor and an O2 microsensor were used to measure profiles across roots of chickpea seedlings in aerated or hypoxic conditions. Simultaneous, nondestructive flux measurements of O2 consumption, CO2 production, and thus RQ, were taken for roots with declining O2 . Radial profiling revealed severe hypoxia and c. 0.8 kPa CO2 within the root vascular cylinder. The distance penetrated by O2 into the roots was shorter at lower O2 . The gradient in CO2 was in the opposite direction to that of O2 , across the roots and diffusive boundary layer. RQ increased as external O2 was lowered. For chickpea roots in solution at air equilibrium, O2 was very low and CO2 was elevated within the vascular cylinder; the extent of the severely hypoxic core increased as external O2 was reduced. The increased RQ in roots in response to declining external O2 highlighted the shift from respiration to ethanolic fermentation as the severely hypoxic/anoxic core became a progressively greater proportion of the root tissues.

Changes in Expression Level of OsHKT1;5 Alters Activity of Membrane Transporters Involved in K+ and Ca2+ Acquisition and Homeostasis in Salinized Rice Roots.

In rice, the OsHKT1;5 gene has been reported to be a critical determinant of salt tolerance. This gene is harbored by the SKC1 locus, and its role was attributed to Na+ unloading from the xylem. No direct evidence, however, was provided in previous studies. Also, the reported function of SKC1 on the loading and delivery of K+ to the shoot remains to be explained. In this work, we used an electrophysiological approach to compare the kinetics of Na+ uptake by root xylem parenchyma cells using wild type (WT) and NIL(SKC1) plants. Our data showed that Na+ reabsorption was observed in WT, but not NIL(SKC1) plants, thus questioning the functional role of HKT1;5 as a transporter operating in the direct Na+ removal from the xylem. Instead, changes in the expression level of HKT1;5 altered the activity of membrane transporters involved in K+ and Ca2+ acquisition and homeostasis in the rice epidermis and stele, explaining the observed phenotype. We conclude that the role of HKT1;5 in plant salinity tolerance cannot be attributed to merely reducing Na+ concentration in the xylem sap but triggers a complex feedback regulation of activities of other transporters involved in the maintenance of plant ionic homeostasis and signaling under stress conditions.

Rhizosecretion of stele-synthesized glucosinolates and their catabolites requires GTR-mediated import in Arabidopsis.

Casparian strip-generated apoplastic barriers not only control the radial flow of both water and ions but may also constitute a hindrance for the rhizosecretion of stele-synthesized phytochemicals. Here, we establish root-synthesized glucosinolates (GLS) are in Arabidopsis as a model to study the transport routes of plant-derived metabolites from the site of synthesis to the rhizosphere. Analysing the expression of GLS synthetic genes in the root indicate that the stele is the major site for the synthesis of aliphatic GLS, whereas indole GLS can be synthesized in both the stele and the cortex. Sampling root exudates from the wild type and the double mutant of the GLS importers GTR1 and GTR2 show that GTR-mediated retention of stele-synthesized GLS is a prerequisite for the exudation of both intact GLS and their catabolites into the rhizosphere. The expression of the GTRs inside the stele, combined with the previous observation that GLS are exported from biosynthetic cells, suggest three possible routes of stele-synthesized aliphatic GLS after their synthesis: (i) GTR-dependent import to cells symplastically connected to the cortical cells and the rhizosphere; (ii) GTR-independent transport via the xylem to the shoot; and (iii) GTR-dependent import to GLS-degrading myrosin cells at the cortex. The study suggests a previously undiscovered role of the import process in the rhizosecretion of root-synthesized phytochemicals.

Revealing the roles of GORK channels and NADPH oxidase in acclimation to hypoxia in Arabidopsis.

Regulation of root cell K+ is essential for acclimation to low oxygen stress. The potential roles of GORK (depolarization-activated guard cell outward-rectifying potassium) channels and RBOHD (respiratory burst oxidase homologue D) in plant adaptive responses to hypoxia were investigated in the context of tissue specificity (epidermis versus stele; elongation versus mature zone) in roots of Arabidopsis. The expression of GORK and RBOHD was down-regulated by 2- to 3-fold within 1 h and 24 h of hypoxia treatment in Arabidopsis wild-type (WT) roots. Interestingly, a loss of the functional GORK channel resulted in a waterlogging-tolerant phenotype, while rbohD knockout was sensitive to waterlogging. To understand their functions under hypoxia stress, we studied K+, Ca2+, and reactive oxygen species (ROS) distribution in various root cell types. gork1-1 plants had better K+ retention ability in both the elongation and mature zone compared with the WT and rbohD under hypoxia. Hypoxia induced a Ca2+ increase in each cell type after 72 h, and the increase was much less pronounced in rbohD than in the WT. In most tissues except the elongation zone in rbohD, the H2O2 concentration had decreased after 1 h of hypoxia, but then increased significantly after 24 h of hypoxia in each zone and tissue, further suggesting that RBOHD may shape hypoxia-specific Ca2+ signatures via the modulation of apoplastic H2O2 production. Taken together, our data suggest that plants lacking functional GORK channels are more capable of retaining K+ for their better performance under hypoxia, and that RBOHD is crucial in hypoxia-induced Ca2+ signalling for stress sensing and acclimation mechanism.

Tissue-specific root ion profiling reveals essential roles of the CAX and ACA calcium transport systems in response to hypoxia in Arabidopsis.

Waterlogging is a major abiotic stress that limits the growth of plants. The crucial role of Ca(2+) as a second messenger in response to abiotic and biotic stimuli has been widely recognized in plants. However, the physiological and molecular mechanisms of Ca(2+) distribution within specific cell types in different root zones under hypoxia is poorly understood. In this work, whole-plant physiological and tissue-specific Ca(2+) changes were studied using several ACA (Ca(2+)-ATPase) and CAX (Ca(2+)/proton exchanger) knock-out Arabidopsis mutants subjected to waterlogging treatment. In the wild-type (WT) plants, several days of hypoxia decreased the expression of ACA8, CAX4, and CAX11 by 33% and 50% compared with the control. The hypoxic treatment also resulted in an up to 11-fold tissue-dependent increase in Ca(2+) accumulation in root tissues as revealed by confocal microscopy. The increase was much higher in stelar cells in the mature zone of Arabidopsis mutants with loss of function for ACA8, ACA11, CAX4, and CAX11 In addition, a significantly increased Ca(2+) concentration was found in the cytosol of stelar cells in the mature zone after hypoxic treatment. Three weeks of waterlogging resulted in dramatic loss of shoot biomass in cax11 plants (67% loss in shoot dry weight), while in the WT and other transport mutants this decline was only 14-22%. These results were also consistent with a decline in leaf chlorophyll fluorescence (F v/F m). It is suggested that CAX11 plays a key role in maintaining cytosolic Ca(2+) homeostasis and/or signalling in root cells under hypoxic conditions.

CLE6 expression recovers gibberellin deficiency to promote shoot growth in Arabidopsis.

Small peptides act as local signals during plant development, but few studies have examined their interaction with phytohormone signaling. Here, we show that application of gibberellin (GA) to Arabidopsis shoots induces substantial accumulation of transcripts encoded by CLE6, a member of the CLAVATA/ESR-RELATED (CLE) gene family, in the root stele, followed by promotion of organ growth by CLE6 in GA-deficient plants. The long-distance effect of GA4 was demonstrated by the observation that its application to the shoot apex of the GA-deficient mutant ga3ox1/ga3ox2 rescued the short-root phenotype. Microarray analysis was used to identify root-expressed genes that respond to systemic application of GA, and CLE6 was selected for further analysis. CLE6 was highly expressed in roots at the young seedling stage, and CLE6 promoter activity was strong in hypocotyls and roots, especially in root stele cells at branch points. Application of CLE6 peptide had no obvious effect on the growth and development of GA-deficient mutant plants. Nonetheless, the fact that ectopic over-expression of CLE6 in the GA-deficient mutant promoted root growth and branching, petiole elongation, bolting rate and stem length showed that CLE6 expression partially compensates for the GA deficiency. Reciprocal grafting of GA-deficient mutant plants to 35S::CLE6 transformants complemented the shoot phenotype associated with GA deficiency, demonstrating the systemic effect of CLE6 from root to shoot. These data suggest that root-expressed CLE6 is systemically involved in shoot growth under GA action in Arabidopsis.

Oxygen deficiency and salinity affect cell-specific ion concentrations in adventitious roots of barley (Hordeum vulgare).

Oxygen deficiency associated with soil waterlogging adversely impacts root respiration and nutrient acquisition. We investigated the effects of O2 deficiency and salinity (100 mM NaCl) on radial O2 concentrations and cell-specific ion distributions in adventitious roots of barley (Hordeum vulgare). Microelectrode profiling measured O2 concentrations across roots in aerated, aerated saline, stagnant or stagnant saline media. X-ray microanalysis at two positions behind the apex determined the cell-specific elemental concentrations of potassium (K), sodium (Na) and chloride (Cl) across roots. Severe O2 deficiency occurred in the stele and apical regions of roots in stagnant solutions. O2 deficiency in the stele reduced the concentrations of K, Na and Cl in the pericycle and xylem parenchyma cells at the subapical region. Near the root apex, Na declined across the cortex in roots from the aerated saline solution but was relatively high in all cell types in roots from the stagnant saline solution. Oxygen deficiency has a substantial impact on cellular ion concentrations in roots. Both pericycle and xylem parenchyma cells are involved in energy-dependent K loading into the xylem and in controlling radial Na and Cl transport. At root tips, accumulation of Na in the outer cell layers likely contributed to reduction of Na in inner cells of the tips.

Overexpression of phosphatidylserine synthase IbPSS1 enhances salt tolerance by stimulating ethylene signaling-dependent lignin synthesis in sweetpotato roots.

Phosphatidylserine (PS) is an important lipid signaling required for plant growth regulation and salt stress adaptation. However, how PS positively regulate plant salt tolerance is still largely unknown. In this study, IbPSS1-overexpressed sweetpotato plants that exhibited overproduction of PS was employed to explore the mechanisms underlying the PS stimulation of plant salt tolerance. The results revealed that the IbPSS1-overexpressed sweetpotato accumulated less Na+ in the stem and leaf tissues compared with the wild type plants. Proteomic profile of roots showed that lignin synthesis-related proteins over-accumulated in IbPSS1-overexpressed sweetpotato. Correspondingly, the lignin content was enhanced but the influx of Na + into the stele was significantly blocked in IbPSS1-overexpressed sweetpotato. The results further revealed that ethylene synthesis and signaling related genes were upregulated in IbPSS1-overexpressed sweetpotato. Ethylene imaging experiment revealed the enhancement of ethylene mainly localized in the root stele. Inhibition of ethylene synthesis completely reversed the PS-overproduction induced lignin synthesis and Na+ influx pattern in stele tissues. Taken together, our findings demonstrate a mechanism by which PS regulates ethylene signaling and lignin synthesis in the root stele, thus helping sweetpotato plants to block the loading of Na+ into the xylem and to minimize the accumulation of Na+ in the shoots.

The worldwide allometric relationship in anatomical structures for plant roots.

The cortex (i.e., absorptive tissue) and stele (transportive vascular tissue) are fundamental to the function of plant roots. Unraveling how these anatomical structures are assembled in absorptive roots is essential for our understanding of plant ecology, physiology, and plant responses to global environmental changes. In this review, we first compile a large data set on anatomical traits in absorptive roots, including cortex thickness and stele radius, across 698 observations and 512 species. Using this data set, we reveal a common root allometry in absorptive root structures, i.e., cortex thickness increases much faster than stele radius with increasing root diameter (hereafter, root allometry). Root allometry is further validated within and across plant growth forms (woody, grass, and liana species), mycorrhiza types (arbuscular mycorrhiza, ectomycorrhiza, and orchid mycorrhizas), phylogenetic gradients (from ferns to Orchidaceae), and environmental change scenarios (e.g., elevation of atmospheric CO2 concentration and nitrogen fertilization). These findings indicate that root allometry is common in plants. Importantly, root allometry varies greatly across species. We then summarize recent research on the mechanisms of root allometry and potential issues regarding these mechanisms. We further discuss ecological and evolutionary implications of root allometry. Finally, we propose several important research directions that should be pursued regarding root allometry.

The primary root procambium contributes to lateral root formation through its impact on xylem connection.

The postembryonic formation of lateral roots (LRs) starts in internal root tissue, the pericycle. An important question of LR development is how the connection of the primary root vasculature with that of the emerging LR is established and whether the pericycle and/or other cell types direct this process. Here, using clonal analysis and time-lapse experiments, we show that both the procambium and pericycle of the primary root (PR) affect the LR vascular connectivity in a coordinated manner. We show that during LR formation, procambial derivates switch their identity and become precursors of xylem cells. These cells, together with the pericycle-origin xylem, participate in the formation of what we call a "xylem bridge" (XB), which establishes the xylem connection between the PR and the nascent LR. If the parental protoxylem cell fails to differentiate, XB is still sometimes formed but via a connection with metaxylem cells, highlighting that this process has some plasticity. Using mutant analyses, we show that the early specification of XB cells is determined by CLASS III HOMEODOMAIN-LEUCINE ZIPPER (HD-ZIP III) transcription factors (TFs). Subsequent XB cell differentiation is marked by the deposition of secondary cell walls (SCWs) in spiral and reticulate/scalariform patterns, which is dependent on the VASCULAR-RELATED NAC-DOMAIN (VND) TFs. XB elements were also observed in Solanum lycopersicum, suggesting that this mechanism may be more widely conserved in plants. Together, our results suggest that plants maintain vascular procambium activity, which safeguards the functionality of newly established lateral organs by assuring the continuity of the xylem strands throughout the root system.

Spatial O2 Profile in Coix lacryma-jobi and Sorghum bicolor along the Gas Diffusion Pathway under Waterlogging Conditions.

While internal aeration in plants is critical for adaptation to waterlogging, there is a gap in understanding the differences in oxygen diffusion gradients from shoots to roots between hypoxia-tolerant and -sensitive species. This study aims to elucidate the differences in tissue oxygen concentration at various locations on the shoot and root between a hypoxia-tolerant species and a -sensitive species using a microneedle sensor that allows for spatial oxygen profiling. Job's tears, a hypoxia-tolerant species, and sorghum, a hypoxia-susceptible species, were tested. Plants aged 10 days were acclimated to a hypoxic agar solution for 12 days. Oxygen was profiled near the root tip, root base, root shoot junction, stem, and leaf. An anatomical analysis was also performed on the roots used for the O2 profile. The oxygen partial pressure (pO2) values at the root base and tip of sorghum were significantly lower than that of the root of Job's tears. At the base of the root of Job's tears, pO2 rapidly decreased from the root cortex to the surface, indicating a function to inhibit oxygen leakage. No significant differences in pO2 between the species were identified in the shoot part. The root cortex to stele ratio was significantly higher from the root tip to the base in Job's tears compared to sorghum. The pO2 gradient began to differ greatly at the root shoot junction and root base longitudinally, and between the cortex and stele radially, between Job's tears and sorghum. Differences in the root oxygen retention capacity and the cortex to stele ratio are considered to be related to differences in pO2.