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ROOT MERISTEM GROWTH FACTOR1 (RGF1) and its receptors RGF1 INSENSITIVEs (RGIs), a group of leucine-rich repeat receptor kinases, promote primary root meristem activity via a mitogen-activated protein kinase (MPK) signaling cascade and control root gravitropism in Arabidopsis. Genetic analyses and in vitro binding assays have indicated that among five RGIs identified in Arabidopsis, RGI1, RGI2, and RGI3 recognize RGF1 peptides. However, it remains unclear whether the RGF1 peptide is redundantly recognized by these RGIs or mainly by a single RGI in the regulation of primary root meristem activity. In the present study, we analyzed root meristem growth of the rgi1 , rgi2 , and rgi3 single mutants in response to RGF1 treatment and observed a significantly decreased sensitivity in meristem growth of rgi1 and complete insensitivity in rgi1 rgi2 rgi3 triple mutant compared with the wild type but not in the rgi1 and rgi2 single mutants. We also observed that both root gravitropism and meristem growth in the BRASSINOSTEROID INSENSITIVE1-ASSOCIATED RECEPTOR KINASE 1 ( bak1 ) single mutant were insensitive to RGF1 peptide treatment, whereas other serk mutants, such as serk1 , serk2 , and serk4 , were fully sensitive to RGF1 peptide like the wild type. These mutant analyses suggest that RGI1-BAK1 pair acts as the main receptor-coreceptor pair for regulating primary root gravitropism and meristem activity in response to RGF1 peptide in Arabidopsis.

Rice (Oryza sativa) production consumes a huge amount of fresh water, and improvement of drought tolerance in rice is important to conserve water resources and minimize yield loss under drought. However, processes to improve drought tolerance in rice have not been fully explored, and a comparative study between rice and wheat (Triticum aestivum) is an effective method to understand the mechanisms determining drought tolerance capacity. In the present study, we applied short-term drought stress to Shanyou 63 rice and Yannong 19 wheat to create a range of water potentials and investigated the responses of gas exchange, plant hydraulic conductance, and root morphological and anatomical traits to soil drought. We found that photosynthesis in rice was more sensitive to drought stress than that in wheat, which was related to differences in the decline of stomatal conductance and plant hydraulic conductance (Kplant). The decline of Kplant under drought was mainly driven by the decrease of soil-root interface hydraulic conductance (Ki) because Ki was more sensitive to drought than root and shoot hydraulic conductance and the soil-root interface contributed to >40% of whole-plant hydraulic resistance in both crops. Root shrinkage in response to drought was more severe in rice than that in wheat, which explains the larger depression of Ki and Kplant under drought stress in rice. We concluded that the decline of Ki drives the depression of Kplant and photosynthesis in both crops, and the plasticity of root morphology and anatomy is important in determining drought tolerance capacity., Competing Interests: Conflict of interest statement. None declared., (© American Society of Plant Biologists 2023. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Microbiota benefit their hosts by improving nutrient uptake and pathogen protection. How host immunity restricts microbiota while avoiding autoimmunity is poorly understood. Here we show that the Arabidopsis phytosulfokine receptor 1 (pskr1) mutant displays autoimmunity (plant stunting, defence-gene expression and reduced rhizosphere bacterial growth) in response to growth-promoting Pseudomonas fluorescens. Microbiome profiling and microbiota colonization showed that PSKR1-mediated reduction in bacterial growth and stunting is largely specific to Pseudomonas. Transcriptional profiling demonstrated that PSKR1 regulates the growth-defence trade-off during Pseudomonas colonization: PSKR1 upregulates plant photosynthesis and root growth but suppresses salicylic-acid-mediated defences. Genetic epistasis experiments showed that pskr1 stunting and restriction of bacterial growth are salicylic acid dependent. Finally, we showed that Pseudomonas, but not other bacteria, induces PSKR1 expression in roots, suggesting that Pseudomonas might manipulate plant signalling to promote its colonization. Our data demonstrate a genetic mechanism to coordinate beneficial functions of the microbiome while preventing autoimmunity., (© 2023. The Author(s), under exclusive licence to Springer Nature Limited.)

Traits of leaves and fine roots are expected to predict the responses and adaptation of plants to their environments. Whether and how fine-root traits (FRTs) are associated with the allocation of foliar phosphorus (P) fractions of desert species in water- and P-poor environments, however, remains unclear. We exposed seedlings of Alhagi sparsifolia Shap. (hereafter Alhagi) treated with two water and four P-supply levels for three years in open-air pot experiments and measured the concentrations of foliar P fractions, foliar traits, and FRTs. The allocation proportion of foliar nucleic acid-P and acid phosphatase (APase) activity of fine roots were significantly higher by 45.94 and 53.3% in drought and no-P treatments relative to well-watered and high-P treatments, whereas foliar metabolic-P and structural-P were significantly lower by 3.70 and 5.26%. Allocation proportions of foliar structural-P and residual-P were positively correlated with fine-root P (FRP) concentration, but nucleic acid-P concentration was negatively correlated with FRP concentration. A tradeoff was found between the allocation proportion to all foliar P fractions relative to the FRP concentration, fine-root APase activity, and amounts of carboxylates, followed by fine-root morphological traits. The requirement for a link between the aboveground and underground tissues of Alhagi was generally higher in the drought than the well-watered treatment. Altering FRTs and the allocation of P to foliar nucleic acid-P were two coupled strategies of Alhagi under conditions of drought and/or low-P. These results advance our understanding of the strategies for allocating foliar P by mediating FRTs in drought and P-poor environments., (© 2023 Scandinavian Plant Physiology Society.)

Magnesium (Mg) and calcium (Ca) are two essential macronutrients in plants; however, the characteristics of Mg and Ca concentrations in organ, subcellular and chemical forms and their relationships in citrus plants, especially under varying Mg supply, are not well understood. In this study, Citrus sinensis seedlings (cv. Xuegan) were cultivated in conditions of Mg deficiency (0 mmol Mg 2+ L -1 ) and Mg sufficiency (2 mmol Mg 2+ L -1 ) to investigate the responses of Mg and Ca homeostasis in different organs and fractions. Compared with Mg sufficiency, Mg deficiency significantly decreased root and shoot growth, with the shoot biomass reduction of branch organs was greater than that of parent organs. In addition to increasing the Ca concentration in the parent stem and lateral root organs, Mg deficiency significantly decreased the concentrations and accumulations of Mg and Ca in citrus seedlings, further altering their distribution in different organs. More than 50% of Ca and Mg were sequestrated in the cell wall and soluble fractions, respectively, with Mg concentration decreasing by 15.4% in roots and 46.9% in leaves under Mg deficiency, while Ca concentration decreased by 27.6% in roots and increased by 23.6% in parent leaves. Approximately 90% of Mg exists in inorganic, water-soluble, and pectate and protein-bound forms, and nearly 90% of Ca exists in water-soluble, pectate and protein-bound, phosphate and oxalate acid forms. Except for the decreased inorganic Mg in roots and water-soluble Mg and Ca in leaves, Mg deficiency increased the proportions of Mg and Ca in all chemical forms. However, Mg deficiency generally increased the Ca/Mg ratio in various organs, subcellular and chemical forms, with negative relationships between Mg concentration and Ca/Mg ratio, and the variations of Mg and Ca were highly separated between Mg supply and organs. In conclusion, our results provide insights into the effects of Mg supply on Mg and Ca homeostasis in citrus plants., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Masson SAS. All rights reserved.)

Aluminium (Al) toxicity is one of the major constraint for crop production in acidic soil, and the inappropriate utilization of nitrogen fertilizer can accelerate soil acidification. Despite previous studies investigating the regulation of nitrogen forms in Al toxicity of plants, the underlying mechanism, particularly at the molecular level, remains unclear. This study aims to uncover the potentially regulatory mechanism of nitrate (NO 3 - ) in the Al resistance of maize and Arabidopsis. NO 3 - conservatively improves Al resistance in maize and Arabidopsis, with nitrate-elevated citrate synthesis and exudation potentially playing critical roles in excluding Al from the root symplast. ZmSLAH2 in maize and AtSLAH1 in Arabidopsis are essential for the regulation of citrate exudation and NO 3 - -promoted Al resistance, with ZmMYB81 directly targeting the ZmSLAH2 promoter to activate its activity. Additionally, NO 3 - transport is necessary for NO 3 - -promoted Al resistance, with ZmNRT1.1A and AtNRT1.1 potentially playing vital roles. The suppression of NO 3 - transport in roots by ammonium (NH 4 + ) may inhibit NO 3 - -promoted Al resistance. This study provides novel insights into the understanding of the crucial role of NO 3 - -mediated signalling in the Al resistance of plants and offers guidance for nitrogen fertilization on acid soils., (© 2023 John Wiley & Sons Ltd.)

Microplastics can affect their surroundings physically and chemically, resulting in diverse effects on plant-soil systems. Similar to other substances (e.g. nutrients and water), microplastics in the environment occur in patches. Such heterogeneous distributions could affect plant responses to plastic pollution. Yet, this has remained untested. We conducted a multispecies experiment including 29 herbaceous plant species and three different microplastic treatments (a control without microplastics, a homogeneous and a heterogeneous microplastic distribution). Based on biomass and root-morphological traits, we assessed how different plastic distributions affect the performance and root-foraging behavior of plants, and whether stronger root foraging is beneficial when microplastics are distributed patchily. Next to general effects on plant productivity and root morphology, we found very strong evidence for root-foraging responses to patchy plastic distributions, with a clear preference for plastic-free patches, resulting in 25% longer roots and 20% more root biomass in the plastic-free patches. Interestingly, however, these foraging responses were correlated with a reduced plant performance, indicating that the benefits of plastic avoidance did not compensate for the associated investments. Our results provide new insights in plant-microplastic interactions and suggest that plants might not just be passively affected by but could also actively respond to environmental plastic pollution., (© 2023. The Author(s).)

Many tree species have developed extensive root systems that allow them to survive in arid environments by obtaining water from a large soil volume. These root systems can transport and redistribute soil water during drought by hydraulic redistribution (HR). A recent study revealed the phenomenon of evaporation-driven hydraulic redistribution (EDHR), which is driven by evaporative demand (transpiration). In this study, we confirmed the occurrence of EDHR in Chinese white poplar (Populus tomentosa) through root sap flow measurements. We utilized microcomputed tomography technology to reconstruct the xylem network of woody lateral roots and proposed conceptual models to verify EDHR from a physical perspective. Our results indicated that EDHR is driven by the internal water potential gradient within the plant xylem network, which requires 3 conditions: high evaporative demand, soil water potential gradient, and special xylem structure of the root junction. The simulations demonstrated that during periods of extreme drought, EDHR could replenish water to dry roots and improve root water potential up to 38.9% to 41.6%. This highlights the crucial eco-physiological importance of EDHR in drought tolerance. Our proposed models provide insights into the complex structure of root junctions and their impact on water movement, thus enhancing our understanding of the relationship between xylem structure and plant hydraulics., Competing Interests: Conflict of interest statement. The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (© American Society of Plant Biologists 2023. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Due to their unique properties, nanomaterials behave peculiarly in biosystems. Regarding plants, the interactions of nanomaterials can be interpreted on a spatial scale: from local interactions in cells to systemic effects on whole plants and on ecosystems. Interpreted on a time scale, the effects of nanomaterials on plants may be immediate or subsequent. At the cellular level, the composition and structure of the cell wall and membranes are modified by nanomaterials, promoting internalization. The effects of nanomaterials on germination and seedling physiology and on the primary and secondary metabolism in the shoot are realized at organ and organism levels. Nanomaterials interact with the beneficial ecological partners of plants. The effects of nanomaterials on plant growth-promoting rhizobacteria and legume-rhizobia symbiosis can be stimulating or inhibitory, depending on the concentration and type of nanomaterial. Nanomaterials exert a negative effect on arbuscular mycorrhiza, and vice versa. Pollinators are exposed to nanomaterials, which may affect plant reproduction. The substances released by the roots influence the availability of nanomaterials in the rhizosphere, and components of plant cells trigger internalization, translocation, and transformation of nanomaterials. Understanding of the multilevel and bidirectional relationship between plants and nanomaterials is of great relevance., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Organ loss occurs frequently during plant and animal evolution. Sometimes, non-functional organs are retained through evolution. Vestigial organs are defined as genetically determined structures that have lost their ancestral (or salient) function. 1 , 2 , 3 Duckweeds, an aquatic monocot family, exhibit both these characteristics. They possess a uniquely simple body plan, variably across five genera, two of which are rootless. Due to the existence of closely related species with a wide diversity in rooting strategies, duckweed roots represent a powerful system for investigating vestigiality. To explore this, we employed a panel of physiological, ionomic, and transcriptomic analyses, with the main goal of elucidating the extent of vestigiality in duckweed roots. We uncovered a progressive reduction in root anatomy as genera diverge and revealed that the root has lost its salient ancestral function as an organ required for supplying nutrients to the plant. Accompanying this, nutrient transporter expression patterns have lost the stereotypical root biased localization observed in other plant species. While other examples of organ loss such as limbs in reptiles 4 or eyes in cavefish 5 frequently display a binary of presence/absence, duckweeds provide a unique snapshot of an organ with varying degrees of vestigialization in closely related neighbors and thus provide a unique resource for exploration of how organs behave at different stages along the process of loss., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2023 The Author(s). Published by Elsevier Inc. All rights reserved.)

Salinity stress is the second most devastating abiotic factor limiting plant growth and yields. Climate changes have significantly increased salinity levels of soil. Besides improving the physiological responses under stress conditions, jasmonates modulate Mycorrhiza-Plant relationships. The present study aimed to evaluate the effects of methyl jasmonate (MeJ) and Funneliformis mosseae (Arbuscular mycorrhizal (AM) on morphology and improving antioxidant mechanisms in Crocus sativus L. under salinity stress. After inoculation with AM, pre-treated C. sativus corms with MeJ were grown under low, moderate, and severe salinity stress. Intense salinity levels damaged the corm, root, total leaf dry weight, and area. Salinities up to 50 mM increased Proline content and Polyphenol oxidase (PPO) activity, but MeJ increased this trend in proline. Generally, MeJ increased anthocyanins, total soluble sugars, and PPO. Total chlorophyll and superoxide dismutase (SOD) activity increased by salinity. The maximum catalase and SOD activities in + MeJ + AM were 50 and 125 mM, respectively, and the maximum total chlorophyll in -MeJ + AM treatment was 75 mM. Although 20 and 50 mM increased plant growth, using mycorrhiza and jasmonate enhanced this trend. Moreover, these treatments reduced the damage of 75 and 100 mM salinity stress. Using MeJ and AM can improve the growth of saffron under various ranges of salinity stress levels; however, in severe levels like 120 mM, this phytohormone and F. mosseae effects on saffron could be adverse., (© 2023. The Author(s).)

This review focuses on the regulation of root water uptake in plants which are exposed to salt stress. Root water uptake is not considered in isolation but is viewed in the context of other potential tolerance mechanisms of plants-tolerance mechanisms which relate to water relations and gas exchange. Plants spend between one third and half of their lives in the dark, and salt stress does not stop with sunset, nor does it start with sunrise. Surprisingly, how plants deal with salt stress during the dark has received hardly any attention, yet any growth response to salt stress over days, weeks, months and years is the integrative result of how plants perform during numerous, consecutive day/night cycles. As we will show, dealing with salt stress during the night is a prerequisite to coping with salt stress during the day. We hope to highlight with this review not so much what we know, but what we do not know; and this relates often to some rather basic questions.

Unravelling belowground strategies is critical for understanding species coexistence and successional dynamics; yet, our knowledge of nutrient acquisition strategies of forest species at different successional stages remains limited. We measured morphological (diameter, specific root length, and root tissue density), architectural (branching ratio), physiological (ammonium, nitrate, and glycine uptake rates) root traits, and mycorrhizal colonisation rates of eight coexisting woody species in an early successional plantation forest in subtropical China. By incorporating physiological uptake efficiency, we revealed a bi-dimensional root economics space comprising of an 'amount-efficiency' dimension represented by morphological and physiological traits, and a 'self-symbiosis' dimension dominated by architectural and mycorrhizal traits. The early pioneer species relied on root-fungal symbiosis, developing densely branched roots with high mycorrhizal colonisation rates for foraging mobile soil nitrate. The late pioneer species invested in roots themselves and allocated effort towards improving uptake efficiency of less-mobile ammonium. Within the root economics space, the covariation of axes with soil phosphorus availability also distinguished the strategy preference of the two successional groups. These results demonstrate the importance of incorporating physiological uptake efficiency into root economics space, and reveal a trade-off between expanding soil physical space exploration and improving physiological uptake efficiency for successional species coexistence in forests., (© 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation.)

Low-temperature stress alters root system architecture. In particular, changes in the levels and response to auxin and cytokinin determine the fate of root architecture and function under stress because of their vital roles in regulating root cell division, differentiation, and elongation. An intricate nexus of genes encoding components of auxin and cytokinin biosynthesis, signaling, and transport components operate to counteract stress and facilitate optimum development. We review the role of auxin transport and signaling and its regulation by cytokinin during root development and stem cell maintenance under low-temperature stress. We highlight intricate mechanisms operating in root stem cells to minimize DNA damage by altering phytohormone levels, and discuss a working model for cytokinin in low-temperatures stress response., Competing Interests: Declaration of interests The authors declare no conflicts of interest., (Copyright © 2022 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

Excess water can induce flooding stress resulting in yield loss, even in wetland crops such as rice (Oryza). However, traits from species of wild Oryza have already been used to improve tolerance to abiotic stress in cultivated rice. This study aimed to establish root responses to sudden soil flooding among eight wild relatives of rice with different habitat preferences benchmarked against three genotypes of O. sativa. Plants were raised hydroponically, mimicking drained or flooded soils, to assess the plasticity of adventitious roots. Traits included were apparent permeance (PA) to O2 of the outer part of the roots, radial water loss, tissue porosity, apoplastic barriers in the exodermis, and root anatomical traits. These were analysed using a plasticity index and hierarchical clustering based on principal component analysis. For example, O. brachyantha, a wetland species, possessed very low tissue porosity compared with other wetland species, whereas dryland species O. latifolia and O. granulata exhibited significantly lower plasticity compared with wetland species and clustered in their own group. Most species clustered according to growing conditions based on PA, radial water loss, root porosity, and key anatomical traits, indicating strong anatomical and physiological responses to sudden soil flooding., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Although environmental factors affecting adventitious root (AR) formation have been examined, how nutrient status affects ARs under waterlogging conditions remains unclear. In this study, plants' performance in responding to AR regulation based on nutrient supply was investigated in terms of plant morphology, physiology and AR traits. Results indicated that Cleistocalyx operculatus possesses higher waterlogging tolerance than Syzygium cumini according to the waterlogging tolerance coefficient, mainly because of the higher fresh weight, porosity and length of AR in C. operculatus. Nutrient supply treatment under a waterlogging condition significantly decreased the fresh weight, length, number, porosity, cortex area of AR and the ratio of cortex-to-stele area in both species relative to those in the waterlogging treatment, but significantly increased the activities and stele areas of AR, and leaf nutrient content. This result showed that nutrient supply caused variations in the morphological and anatomical structures of AR that were more beneficial to improve nutrient transportation than oxygen absorption under waterlogging conditions, supporting the nutrient-priority hypothesis. Moreover, nutrient supply under waterlogging conditions induced greater increase in stele area of ARs, fresh weight of the whole plant, total leaf area, leaf nitrogen level, total chlorophyll content, net photosynthesis rate and maximum photochemical quantum yield of PSII in S. cumini than in C. operculatus, suggesting that S. cumini can transport more nutrients and easily adapts to increase in nutrient supply under waterlogging conditions. Thus, S. cumini have better performance in extracting and utilizing nutrients in the water for plant growth. The findings showed that terrestrial arbor plants have physiological and microstructural mechanisms that respond to nutrient supply under waterlogging conditions and provide novel insights into the phytoremediation of eutrophic water bodies in wetland systems., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

The first symptom of aluminum (Al) toxicity is the inhibition of root growth, which has been associated with low leaf hydration, with negative consequences for leaf gas exchange including stomatal conductance (gs) observed in many plant species. Here we asked whether low leaf hydration occurs before or after the inhibition of root growth of Citrus × limonia Osbeck ('Mandarin' lime) cultivated for 60 days in nutrient solution with 0 and 1480 μM Al. The length, diameter, surface area and biomass of roots of plants exposed to Al were lower than control plants only at 30 days after treatments (DAT). Until the end of the study, estimated gs (measured by sap flow techniques) was lower than in control plants from 3 DAT, total plant transpiration (Eplant) and root hydraulic conductivity (Lpr) at 7 DAT, and midday leaf water potential (Ψmd) and relative leaf water content at 15 DAT. Abscisic acid (ABA) in leaves was twofold higher in Al-exposed plants 1 DAT, and in roots a twofold higher peak was observed at 15 DAT. As ABA in leaves approached values of control plants after 15 DAT, we propose that low gs of plants exposed to Al is primarily caused by ABA, and the maintenance of low gs could be ascribed to the low Lpr from 7 DAT until the end of the study. Therefore, the low leaf hydration in 'Mandarin' lime exposed to Al does not seem to be caused by root growth inhibition or by a simple consequence of low water uptake due to a stunted root system., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Drought stress often leads to heavy losses in mulberry planting, especially for fruits and leaves. Application of plant growth-promoting fungi (PGPF) endows various plant beneficial traits to overcome adverse environmental conditions, but little is known about the effects on mulberry under drought stress. In the present study, we isolated 64 fungi from well-growing mulberry trees surviving periodical drought stress, and Talaromyces sp. GS1, Pseudeurotium sp. GRs12, Penicillium sp. GR19, and Trichoderma sp. GR21 were screened out due to their strong potential in plant growth promotion. Co-cultivation assay revealed that PGPF stimulated mulberry growth, exhibiting increased biomass and length of stems and roots. Exogenous application of PGPF could alter fungal community structures in the rhizosphere soils, wherein Talaromyces was obviously enhanced after inoculation of Talaromyces sp. GS1, and Peziza was increased in the other treatments. Moreover, PGPF could promote iron and phosphorus absorption of mulberry as well. Additionally, the mixed suspensions of PGPF induced the production of catalase, soluble sugar, and chlorophyll, which in turn enhanced the drought tolerance of mulberry and accelerated their growth recovery after drought. Collectively, these findings might provide new insights into improving mulberry drought tolerance and further boosting mulberry fruit yields by exploiting interactions between hosts and PGPF.

Plants have evolved elaborate mechanisms to sense, respond to and overcome the detrimental effects of high soil salinity. The role of calcium transients in salinity stress signaling is well established, but the physiological significance of concurrent salinity-induced changes in cytosolic pH remains largely undefined. Here, we analyzed the response of Arabidopsis roots expressing the genetically encoded ratiometric pH-sensor pHGFP fused to marker proteins for the recruitment of the sensor to the cytosolic side of the tonoplast (pHGFP-VTI11) and the plasma membrane (pHGFP-LTI6b). Salinity elicited a rapid alkalinization of cytosolic pH (pHcyt) in the meristematic and elongation zone of wild-type roots. The pH-shift near the plasma membrane preceded that at the tonoplast. In pH-maps transversal to the root axis, the epidermis and cortex had cells with a more alkaline pHcyt relative to cells in the stele in control conditions. Conversely, seedlings treated with 100 mM NaCl exhibited an increased pHcyt in cells of the vasculature relative to the external layers of the root, and this response occurred in both reporter lines. These pHcyt changes were substantially reduced in mutant roots lacking a functional SOS3/CBL4 protein, suggesting that the operation of the SOS pathway mediated the dynamics of pHcyt in response to salinity.

Haustoria of root-parasitic plants draw nutrients from the roots of host species. While recent studies have assessed host preferences of parasitic plants, how root-exuded chemicals can mediate host tropism and selection by root-parasitic plants is poorly understood. Under greenhouse conditions, we performed two pot experiments to determine whether the root parasite Santalum album selectively forages for superior hosts (N2-fixing Acacia confusa Merr. or Dalbergia odorifera T. Chen) rather than for inferior hosts (non-N2-fixing Bischofia polycarpa (levl.) Airy Shaw or Dracontomelon duperreranum Pierre), and whether S. album uses host root exudates and/or specific chemicals in these root exudates to locate and trigger haustorium formation. Lateral roots and haustoria of S. album seedlings exhibited greater growth in the direction of D. odorifera roots than toward roots from the other three hosts. Comparative metabolic analysis revealed that D. odorifera root exudates were enriched in isoflavonoid, flavonoid and flavone/flavonol biosynthesis pathways, and that the relative contents of flavonoids were significantly greater in the root exudates of D. odorifera than in those of the other three hosts. Root exudates from D. odorifera significantly promoted S. album root growth, haustorium formation and reactive oxygen species accumulation in haustoria. Our results demonstrate that the key step in plant parasitism by S. album is based on root exudation by a host plant; the exudates function as a metabolite signal that activate lateral root growth and haustorium formation. Our results also indicate that flavonoids in the root exudates could play an important role in S. album foraging activity. Information on the responses of root parasites to host root exudates and/or haustorium-inducing chemicals may be useful for selecting superior host species to plant with valuable species of root parasites., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Mulberry is an economically important crop for sericulture in China. Mulberry plantations are shifting inland, where they face high salinity. Arbuscular mycorrhizal fungi (AMF) reportedly enhance mulberry's tolerance to salinity. Here, we assessed if additional adaptive advantages against salinity are provided by sex differences beyond those provided by mycorrhizal symbiosis. In a pot experiment, male and monoecious plants were exposed to three salinity regimes (0, 50, and 200 mM NaCl) and two mycorrhiza-suppressed conditions (with or without benomyl application) for more than 16 months. We noticed that salinity alone significantly decreased the mycorrhizal colonization rate, salinity tolerance, K + concentrations, and the ionic ratios of all plants. Mycorrhizal association mildly ameliorated the salt-induced detrimental effects, especially for monoecious plants, and sex-specific responses were observed. Meanwhile, both sexes had adopted different strategies to enhance their salinity resistance. Briefly, mycorrhizal monoecious plants exhibited a higher net photosynthetic rate and lower translocation of Na + from root to shoot compared with mycorrhizal males under saline conditions. Their salt tolerance was probably due to the Ca 2+ /Na + in roots. In comparison, male plants exhibited lower Na + acquisition, more Na + translocated from root to shoot, higher root biomass allocation, and higher N concentrations under harsh saline conditions, and their salt tolerance was mainly related to the K + /Na + in their shoots. In conclusion, our results highlight that AMF could be a promising candidate for improving plant performance under highest salinity, especially for monoecious plants. Cultivators must be mindful of applying fungicides, such as benomyl, in saline areas., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Masson SAS. All rights reserved.)

Background: The kinetic parameters of peach rootstock are a major factor contributing to its nitrogen (N) uptake efficiency, which directly affects the yield and quality parameters. However, the impact of grafting on N kinetic parameters is not sufficiently known as rootstocks are grafted with other scion cultivars before being transplanted to the field. The objective of this study was therefore to evaluate whether the grafting of 'Okinawa' rootstock with the scion cultivar 'Chimarrita' can alter the kinetic parameters related to N uptake., Results: The plants were acclimatized in Hoagland solution for 21 days and, sequentially, were tested to assess the depletion of the internal reserves at 15 and 30 days in CaSO4 solution. Morphological and physiological parameters were evaluated, and kinetic parameters were calculated. The depletion period for to reach the minimum concentration (C min ) of NO 3 - and NH 4 + in the plant was 62 h and 56 h, respectively. The optimal time for internal reserve depletion for 'Okinawa' rootstock with and without the 'Chimarrita' scion was 30 days., Conclusion: 'Okinawa' rootstock grafted with 'Chimarrita' scion cultivar was more efficient in NO 3 - and NH 4 + uptake. © 2022 Society of Chemical Industry., (© 2022 Society of Chemical Industry.)

Background: Methane (CH 4 ) and brassinosteroids (BRs) are important signaling molecules involved in a variety of biological processes in plants., Results: Here, marigold (Tagetes erecta L. 'Marvel') was used to investigate the role and relationship between CH 4 and BRs during adventitious root (AR) formation. The results showed a dose-dependent effect of CH 4 and BRs on rooting, with the greatest biological effects of methane-rich water (MRW, CH 4 donor) and 2,4-epibrassinolide (EBL) at 20% and 1 μmol L - 1 , respectively. The positive effect of MRW on AR formation was blocked by brassinoazole (Brz, a synthetic inhibitor of EBL), indicating that BRs might be involved in MRW-regulated AR formation. MRW promoted EBL accumulation during rooting by up-regulating the content of campestanol (CN), cathasterone (CT), and castasterone (CS) and the activity of Steroid 5α-reductase (DET2), 22α-hydroxylase (DWF4), and BR-6-oxidase (BR6ox), indicating that CH 4 could induce endogenous brassinolide (BR) production during rooting. Further results showed that MRW and EBL significantly down-regulated the content of cellulose, hemicellulose and lignin during rooting and significantly up-regulated the hydrolase activity, i.e. cmcase, xylanase and laccase. In addition, MRW and EBL also significantly promoted the activity of two major cell wall relaxing factors, xyloglucan endotransglucosylase/hydrolase (XTH) and peroxidase, which in turn promoted AR formation. While, Brz inhibited the role of MRW on these substances., Conclusions: BR might be involved in CH 4 -promoted AR formation by increasing cell wall relaxation., (© 2022. The Author(s).)

Plants have a broad capacity to regenerate damaged organs. The study of wounding in multiple developmental systems has uncovered many of the molecular properties underlying plants' competence for regeneration at the local cellular level. However, in nature, wounding is rarely localized to one place, and plants need to coordinate regeneration responses at multiple tissues with environmental conditions and their physiological state. Here, we review the evidence for systemic signals that regulate regeneration on a plant-wide level. We focus on the role of auxin and sugars as short- and long-range signals in natural wounding contexts and discuss the varied origin of these signals in different regeneration scenarios. Together, this evidence calls for a broader, system-wide view of plant regeneration competence., (© 2022 The Authors New Phytologist © 2022 New Phytologist Foundation.)

Plants will interact with beneficial endophytic fungi to increase resistance under environmental stress. Among these stresses, salt stress poses one of the major threats to plant growth worldwide. We have studied the response mechanism of Chaetomium globosum D5, a salt-tolerant fungus isolated from the roots of Paeonia lactiflora under salt stress, and its mechanism of action in helping P. lactiflora alleviate salt stress. In our study, high levels of salt inhibit growth, whereas low levels promote the growth of C. globosum D5, which resists salt stress by forming dense hyphae and producing more pigments, soluble proteins, and antioxidants. Under salt stress, growth and photosynthesis of P. lactiflora are inhibited, and they are subjected to osmotic stress, oxidative stress, and ionic stress. C. globosum D5 could help P. lactiflora promote growth and photosynthesis by increasing the uptake of nitrogen and phosphorus and increasing the accumulation of the carbon and photosynthetic pigments, help P. lactiflora alleviate osmotic stress by increasing the accumulation of proline, help P. lactiflora alleviate ion stress by reducing Na + and increasing K + /Na + , Ca2 + /Na + and Mg 2+ /Na  +  ratios in P. lactiflora roots and leaves. In summary, joint action between P. lactiflora and C. globosum D5 is responsible for mitigating damage caused by P. lactiflora under salt stress. We first investigate the interaction between the fungus and P. lactiflora under salt stress, providing a theoretical basis for further investigations into the mechanisms of P. lactiflora's response to salt stress and its promotion in coastal areas., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier GmbH. All rights reserved.)

Gravitropism is an important strategy for the adaptation of plants to the changing environment. Previous reports indicated that Ca 2+ participated in plant gravity response. However, present information on the functions of Ca 2+ in plant gravitropism was obtained mainly on coleoptiles, hypocotyls, and petioles, little is known about the dynamic changes of Ca 2+ during root gravitropism. In the present study, the transgenic Arabidopsis thaliana R-GECO1 was placed horizontally and subsequently vertically on a refitted Leica SP8 laser scanning confocal microscopy with a vertical stage. Real-time observations indicated that gravistimulation induced not only an increase in the Ca 2+ concentration, but also an accelerated occurrence of Ca 2+ sparks in the root cap, especially in the lower side of the lateral root cap, indicating a strong tie between Ca 2+ dynamics and gravistimulation during the early stage of root gravity response.

When it comes to root and mycorrhizal associations that define resource acquisition strategy, there is a need to identify the leading dimension across root physiology, morphology, architecture and whole plant biomass allocation to better predict the plant's responses to multiple environmental constraints. Here, we developed a new framework for understanding the variation in roots and symbiotic fungi by quantifying multiple-scale characteristics, ranging from anatomy to the whole plant. We chose the rubber (Hevea brasiliensis) grown at three elevations to test our framework and to identify the key dimensions for resource acquisition. Results showed that the quantities of absorptive roots and root system architecture, rather than single root traits, played the leading role in belowground resource acquisition. As the elevation increased from the low to high elevation, root length growth, productivity and root mass fraction (RMF) increased by 2.9-, 2.3- and 13.8-fold, respectively. The contribution of RMF to the changes in total root length was 3.6-fold that of specific root length (SRL). Root architecture exhibited higher plasticity than anatomy and morphology. Further, mycorrhizal colonization was highly sensitive to rising elevations with a non-monotonic pattern. By contrast, both leaf biomass and specific leaf area (traits) co-varied with increasing elevation. In summary, rubber trees changed root system architecture by allocating more biomass and lowering the reliance on mycorrhizal fungi rather than improving single root efficiency in adapting to high elevation. Our framework is instructive for traits-based ecology; accurate assessments of forest carbon cycling in response to resource gradient should account for the leading dimension of root system architecture., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Gravity is one of the fundamental environmental cues that affect plant development. Indeed, the plant architecture in the shoots and roots is modulated by gravity. Stems grow vertically upward, whereas lateral organs, such as the lateral branches in shoots, tend to grow at a specific angle according to a gravity vector known as the gravitropic setpoint angle (GSA). During this process, gravity is sensed in specialised gravity-sensing cells named statocytes, which convert gravity information into biochemical signals, leading to asymmetric auxin distribution and driving asymmetric cell division/expansion in the organs to achieve gravitropism. As a hypothetical offset mechanism against gravitropism to determine the GSA, the anti-gravitropic offset (AGO) has been proposed. According to this concept, the GSA is a balance of two antagonistic growth components, that is gravitropism and the AGO. Although the nature of the AGO has not been clarified, studies have suggested that gravitropism and the AGO share a common gravity-sensing mechanism in statocytes. This review discusses the molecular mechanisms underlying gravitropism as well as the hypothetical AGO in the control of the GSA., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

Accessing freshwater resources for agriculture becomes more complex due to increasing demands and declining water quality. Alternative water sources, such as saline water, require ad hoc solutions. Therefore, understanding roots' response to saline water is crucial for future agriculture. We examined the response of three grapevine rootstocks (Paulsen 1103, Richter 110 and SO4) to salt stress. The rootstocks were subjected to two salinity treatments: 10 mM and 30 mM NaCl (EC = 2 and 4 ds/m, respectively). Root and shoot samples were taken at the end of the experiment for morphologic and ionomic analyses. The specific root area (SRA) increased in response to salinity for all three rootstocks due to root tissue density and average root diameter reductions. Salinity also led to increased root Na + and Cl - contents and reduced root K + /Na + ratio, parallel to increased leaf Cl - but not Na + contents. SO4 showed improved chloride and sodium exclusion, concomitant with its highest SRA, resulting from the increase in its thin roots' contribution to the total root system surface area. We suggest that enhanced SRA combined with decreased root tissue density and diameter may improve grapevines' salt exclusion by less salt uptake from the soil., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier B.V. All rights reserved.)

Background and Aims: Light competition can induce varying above-ground responses in plants. However, very little is known regarding the effect of above-ground light competition cues on plant responses below ground. Here we asked whether light competition cues that indicate the occurrence and direction of neighbours above ground might affect directional root placemat., Methods: In a common-garden experiment, we examined the integrated responses of the annual procumbent plant Portulaca oleracea to light competition cues and soil nutrient distribution. Soil nutrients were distributed either uniformly or in patches, and light competition was simulated using a transparent green filter, which was spatially located either in the same or opposite direction of the soil nutrient patch., Key Results: As predicted, root proliferation of P. oleracea increased in the direction of the enriched soil patches but was homogenously distributed under the uniform nutrient distribution. Interestingly, root distribution was also affected by the light competition cue and increased in its direction regardless of the location of the soil patches., Conclusions: Our results provide initial support to the idea that below-ground plant responses to competition might also be regulated by above-ground neighbour cues, highlighting the need to further investigate the combined effects of both above- and below-ground competition cues on root behaviour., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Background: The mechanisms of abscisic acid (ABA) and auxin (IAA) in inducing adventitious root (AR) formation, biomass accumulation, and plant development under long-term waterlogging (LT-WL) conditions are largely unexplored. This study aimed to determine the roles of exogenous application of ABA and IAA in two woody plants (Cleistocalyx operculatus and Syzygium jambos) under LT-WL conditions. A pot experiment was conducted using a complete randomized design with two factors: (i) LT-WL and (ii) application of exogenous phytohormones (ABA and IAA) for 120 d., Results: Results revealed that exogenous ABA and IAA promoted LT-WL tolerance in both species. In C. operculatus and S. jambos, plant height, the number of blades, leaf area, and fresh shoot weight were increased by exogenous IAA under LT-WL. However, exogenous ABA affected more the adventitious and primary root in C. operculatus compared to S. jambos. LT-WL decreased drastically the photosynthetic activities in both species, but adding moderate amounts of exogenous ABA or IAA protected the photosynthesis apparatus under LT-WL. Exogenous phytohormones at certain levels decreased the superoxide anion level and malondialdehyde accumulation in plants under LT-WL. Also, the increase of the peroxidases and superoxide dismutase activities by exogenous phytohormones was more marked in C. operculatus compared to S. jambos. Meanwhile, the catalase activity was down-regulated in both species by exogenous phytohormones. Exogenous ABA or IAA positively regulated the jasmonic acid content in ARs under LT-WL. Moderate application of exogenous ABA or IAA in plants under LT-WL decreased the ABA content in the leaves. Lower accumulation of IAA and ABA in the leaves of C. operculatus under LT-WL was positively correlated with a decrease in antioxidant activity., Conclusions: Lastly, C. operculatus which has greater morphology indexes was more tolerant to waterlogging than S. jambos. Moreover, the adaptive strategies via exogenous ABA were more built around the below-ground biomass indexes particularly in C. operculatus, while exogenous IAA backed the above-ground biomass in both species. Overall, the exogenous hormones applied (spraying or watering) influenced differentially the plant's responses to LT-WL. The phytohormonal profile of plants exposed to waterlogging stress varied depending on the species' tolerance level., (© 2022. The Author(s).)

Suberin in roots acts as a physical barrier preventing water/mineral losses. In Arabidopsis, root suberization is regulated by abscisic acid (ABA) and ethylene in response to nutrient stresses. ABA also mediates coordination between microbiota and root endodermis in mineral nutrient homeostasis. However, it is not known whether this regulatory system is common to plants in general, and whether there are other key molecule(s) involved. We show that serotonin acts downstream of ABA in regulating suberization in rice and Arabidopsis and negatively regulates suberization in rice roots in response to salinity. We show that ABA represses transcription of the key gene (OsT5H) in serotonin biosynthesis, thus promoting root suberization in rice. Conversely, overexpression of OsT5H or supplementation with exogenous serotonin represses suberization and reduces tolerance to salt stress. These results identify an ABA-serotonin regulatory module controlling root suberization in rice and Arabidopsis, which is likely to represent a general mechanism as ABA and serotonin are ubiquitous in plants. These findings are of significant importance to breeding novel crop varieties that are resilient to abiotic stresses and developing strategies for production of suberin-rich roots to sequestrate more CO 2 , helping to mitigate the effects of climate change., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

Key Message: The present study clearly showed that the optimum root zone temperature of photosynthesis and plant growth was affected by air temperature, and that optimization of root zone temperature depending on an air growth temperature by cooling systems could lead to improvement of plant production. Temperature is one of the critical factors affecting plant growth and yield production. Both air and root zone temperatures can strongly affect growth and development of plants. However, studies on the effects of root zone temperature on plant growth parameters along with air temperature are still limited. In the present study, the effects of air and root zone temperature on plant growth, physiological parameters and photosynthetic characteristics of lettuce plants were investigated to optimize the air and root zone temperature to achieve the best growth conditions for lettuce plants. Two air temperature treatments (30/25 and 25/20 °C at day/night temperature) and five root zone temperature treatments (15, 20, 25, 30 and 35 °C) were applied in this study. The present study showed that the maximum plant growth of lettuce plants was higher in air temperatures at 30/25 °C than in 25/20 °C. When the plants were grown at an air temperature of 30/25 °C, the optimum root zone temperature appeared to be 30 °C. However, when the plants were grown at an air temperature of 25/20 °C, the optimum root temperature decreased and appeared to be 25 °C. Furthermore, plants grown under air temperature of 30/25 °C showed greater CO 2 assimilation rate, stomatal conductance, electron transport rate (ETR) at high light, and lower non-photochemical quenching (NPQ) at high light than those of 25/20 °C. These results suggest that it is necessary to control and adjust the root zone temperature based on the air temperature., (© 2022. The Author(s), under exclusive licence to Springer Nature B.V.)

In recent years, the effects of plastic contamination on soil and plants have received growing attention. Plastic can affect soil water content and thus may interact with the effects of drought on soil and plants. However, the effects of plastic on soil are highly context-dependent, and interactions with drought have been hardly tested. We conducted two greenhouse experiments to test the combined effects of plastic fragments (of varying size and concentration), water availability and soil texture, on soil water content and performance of the plant Arabidopsis thaliana. Plastic fragments had stronger negative effects on soil water content in low water availability, and the shape of this response (linear vs. unimodal) was mediated by soil texture. Conversely, increasing concentration of plastic had positive effects on plant growth. We suggest that plastic fragments introduce fracture points within soil aggregates. This increases number and size of soil pores favoring water loss but also facilitating root growth. Our results suggest complex interactive effects of plastic and drought, that may lead to a decoupling of plant and soil response. These processes should be taken into account in ecological studies and agricultural practices., (© 2022. The Author(s).)

Background and Aims: Plant performance is enhanced by balancing above- and below-ground resource uptake through the intraspecific adjustment of leaf and root traits. It is assumed that these organ adjustments are at least partly coordinated, so that analogous leaf and root traits broadly covary. Understanding the extent of such intraspecific leaf-root trait covariation would strongly contribute to our understanding of how plants match above- and below-ground resource use strategies as their environment changes, but comprehensive studies are lacking., Methods: We measured analogous leaf and root traits from 11 species, as well as climate, soil and vegetation properties along a 1000-m elevation gradient in the French Alps. We determined how traits varied along the gradient, to what extent this variation was determined by the way different traits respond to environmental cues acting at different spatial scales (i.e. within and between elevations), and whether trait pairs covaried within species., Key Results: Leaf and root trait patterns strongly diverged: across the 11 species along the gradient, intraspecific leaf trait patterns were largely consistent, whereas root trait patterns were highly idiosyncratic. We also observed that, when compared with leaves, intraspecific variation was greater in root traits, due to the strong effects of the local environment (i.e. at the same elevation), while landscape-level effects (i.e. at different elevations) were minor. Overall, intraspecific trait correlations between analogous leaf and root traits were nearly absent., Conclusions: Our study suggests that environmental gradients at the landscape level, as well as local heterogeneity in soil properties, are the drivers of a strong decoupling between analogous leaf and root traits within species. This decoupling of plant resource acquisition strategies highlights how plants can exhibit diverse whole-plant acclimation strategies to modify above- and below-ground resource uptake, improving their resilience to environmental change., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Background: Despite the importance of root hydraulics, there is little research on the in situ dynamic responses of embolism formation and embolism repair of roots distributed in different soil depths in response to different water regimes., Results: The vessel diameter, hydraulic conductivity, and vulnerability to cavitation were in the order of deep root > shallow root > branch. The midday PLC of shallow root was the highest in the dry season, while the midday PLC of deep root slightly higher than that of branch with no significant difference in the two seasons. The capacity of embolism repair of roots was significantly greater than that of branch both in dry season and wet season. The xylem pressure was in the order of deep roots > shallow root > branch, and it was negative in most of the time for the latter two in the dry season, but positive for both of the roots during the observation period in the wet season. The NSC and starch content in roots were significantly higher than those in branches, especially in the dry season. In contrast, roots had lower content of soluble sugar., Conclusions: The relatively stable water condition in soil, especially in the deep layers, is favorable for the development of larger-diameter vessels in root xylem, however it cannot prevent the root from forming embolism. The mechanism of embolism repair may be different in different parts of plants. Deep roots mainly depend on root pressure to refill the embolized vessels, while branches mainly depend on starch hydrolysis to soluble sugars to do the work, with shallow roots shifted between the two mechanisms in different moisture regimes. There is theoretically an obvious trade-off between conducting efficiency and safety over deep roots, shallow roots and branches. But in natural conditions, roots do not necessarily suffer more severe embolism than branches, maybe due to their root pressure-driven embolism repair and relatively good water conditions., (© 2022. The Author(s).)

The efficient uptake of nutrients depends on the ability of roots to respond to gradients of these resources. Although pot experiments have shown that species differ in their ability to proliferate their roots in nutrient-rich patches, the role of such differences in determining root shapes in the field is unclear. We used fine-scale quantitative (q)PCR-based species-specific mapping of roots in a grassland community to reconstruct species-specific root system shapes. We linked them with data from pot experiments on the ability of these species to proliferate in nutrient-rich patches and their rooting depth. We found remarkable diversity in root system shapes, from cylindrical to conical. Interspecific differences in rooting depths in pots were the main determinant of rooting depths in the field, whereas differences in foraging ability played only a minor role. Although some species with strong foraging ability did place their roots into nutrient-rich soil layers, it was not a universal pattern. The results imply that although the vertical differentiation of grassland species is pronounced, it is primarily not driven by the differential plastic response of species to soil nutrient gradients. This may constrain the coexistence of species with similar rooting depths and may instead favour coexistence of species differing in their architectural blueprints., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

Nuclear Ca 2+ oscillations allow symbiosis signaling, facilitating plant recognition of beneficial microsymbionts, nitrogen-fixing rhizobia, and nutrient-capturing arbuscular mycorrhizal fungi. Two classes of channels, DMI1 and CNGC15, in a complex on the nuclear membrane, coordinate symbiotic Ca 2+ oscillations. However, the mechanism of Ca 2+ signature generation is unknown. Here, we demonstrate spontaneous activation of this channel complex, through gain-of-function mutations in DMI1 , leading to spontaneous nuclear Ca 2+ oscillations and spontaneous nodulation, in a CNGC15 -dependent manner. The mutations destabilize a hydrogen-bond or salt-bridge network between two RCK domains, with the resultant structural changes, alongside DMI1 cation permeability, activating the channel complex. This channel complex was reconstituted in human HEK293T cell lines, with the resultant calcium influx enhanced by autoactivated DMI1 and CNGC15s. Our results demonstrate the mode of activation of this nuclear channel complex, show that DMI1 and CNGC15 are sufficient to create oscillatory Ca 2+ signals, and provide insights into its native mode of induction.

Root foraging enables plants to obtain more soil nutrients in a constantly changing nutrient environment. Little is known about the adaptation mechanism of adventitious roots of plants dominated by asexual reproduction (such as tea plants) to soil potassium heterogeneity. We investigated root foraging strategies for K by two tea plants (low-K tolerant genotype "1511" and low-K intolerant genotype "1601") using a multi-layer split-root system. Root exudates, root architecture and transcriptional responses to K heterogeneity were analyzed by HPLC, WinRHIZO and RNA-seq. With the higher leaf K concentrations and K biological utilization indexes, "1511" acclimated to K heterogeneity better than "1601". For "1511", maximum total root length and fine root length proportion appeared on the K-enriched side; the solubilization of soil K reached the maximum on the low-K side, which was consistent with the amount of organic acids released through root exudation. The cellulose decomposition genes that were abundant on the K-enriched side may have promoted root proliferation for "1511". This did not happen in "1601". The low-K tolerant tea genotype "1511" was better at acclimating to K heterogeneity, which was due to a smart root foraging strategy: more roots (especially fine roots) were developed in the K-enriched side; more organic acids were secreted in the low-K side to activate soil K and the root proliferation in the K-enriched side might be due to cellulose decomposition. The present research provides a practical basis for a better understanding of the adaptation strategies of clonal woody plants to soil nutrient availability.

Mangroves are frequently inundated with saline water and have evolved different anatomical and physiological mechanisms to filter and, in some species, excrete excess salt from the water they take up. Because salts impose osmotic stress, interspecific differences in salt tolerance and salt management strategy may influence physiological responses to drought throughout the entire plant hydraulic pathway, from roots to leaves. Here, we characterized embolism vulnerability simultaneously in leaves, stems, and roots of seedlings of two mangrove species (Avicennia marina and Bruguiera gymnorrhiza) along with turgor-loss points in roots and leaves and xylem anatomical traits. In both species, the water potentials causing 50% of total embolism were less negative in roots and leaves than they were in stems, but the water potentials causing incipient embolism (5%) were similar in roots, stems, and leaves. Stomatal closure in leaves and turgor loss in both leaves and roots occurred at water potentials only slightly less negative than the water potentials causing 5% of total embolism. Xylem anatomical traits were unrelated to vulnerability to embolism. Vulnerability segmentation may be important in limiting embolism spread into stems from more vulnerable roots and leaves. Interspecific differences in salt tolerance affected hydraulic traits from roots to leaves: the salt-secretor A. marina lost turgor at more negative water potentials and had more embolism-resistant xylem than the salt-excluder B. gymnorrhiza. Characterizing physiological thresholds of roots may help to explain recent mangrove mortality after drought and extended saltwater inundation., (© American Society of Plant Biologists 2022. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Plant survival during environmental stress greatly affects ecosystem carbon (C) cycling, and plant-microbe interactions are central to plant stress survival. The release of C-rich root exudates is a key mechanism plants use to manage their microbiome, attracting beneficial microbes and/or suppressing harmful microbes to help plants withstand environmental stress. However, a critical knowledge gap is how plants alter root exudate concentration and composition under varying stress levels. In a greenhouse study, we imposed three drought treatments (control, mild, severe) on blue grama (Bouteloua gracilis Kunth Lag. Ex Griffiths), and measured plant physiology and root exudate concentration and composition using GC-MS, NMR, and FTICR. With increasing drought severity, root exudate total C and organic C increased concurrently with declining predawn leaf water potential and photosynthesis. Root exudate composition mirrored the physiological gradient of drought severity treatments. Specific compounds that are known to alter plant drought responses and the rhizosphere microbiome mirrored the drought severity-induced root exudate compositional gradient. Despite reducing C uptake, these plants actively invested C to root exudates with increasing drought severity. Patterns of plant physiology and root exudate concentration and composition co-varied along a gradient of drought severity., (© 2022. The Author(s).)

To understand the regulation of roots plasma membrane H+-ATPase in Masson pine responding to acid deposition, the changes in biomass, plant morphology, intracellular H+, enzyme activity and H+-ATPase genes expression in Masson pine seedlings exposed to simulated acid rain (SAR, pH 5.6 and 4.6) with and without vanadate were studied. Simulated acid rain exposure for 60 days increased the intracellular H+ in pine roots whether added with 0.1 mM Na3VO4 or not. The growth of seedlings treated with SAR was maintained well, even the primary lateral root length, root dry weight and number of root tips in seedlings exposed to SAR at pH 4.6 were higher than that of the control (pH 6.6). However, the addition of vanadate resulted in severe growth inhibition and obvious decline in morphological parameters. Similarly, ATP hydrolytic activity and H+ transport activity of roots plasma membrane H+-ATPase, both were stimulated by SAR whereas they were inhibited by vanadate, and the highest activity stimulation was observed in pine roots subjected to SAR at pH 4.6. In addition, SAR also induced the expression of the investigated H+-ATPase subunits (atpB, atpE, atpF, atpH and atpI). Therefore, the roots plasma membrane H+-ATPase is instrumental in the growth of Masson pine seedlings adapting to acid rain by a manner of pumping more protons across the membrane through enhancing its activity, and which involves the upregulated gene expression of roots H+-ATPase subunits at transcriptional level., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

The variation in fine root respiration with root age provides insight into root adaptation to climate warming, but the mechanism is poorly understood. In this study, we investigated the respiratory response of fine roots (<1 mm and 1-2 mm) of different ages (2-, 4- and 6-month old) of Chinese fir (Cunninghamia lanceolata (Lamb.)) seedlings to soil warming (4 °C above the control using cable heating). Fine roots were excised to measure the specific respiration rate at a reference temperature of 20 °C (SRR20), and root morphological and chemical traits were measured. Soil warming significantly increased SRR20 by 40% compared with the control, potentially indicating limited acclimation on a short time scale (6 months). However, soil warming increased SRR20 significantly in 2-month-old roots (by 72%) compared with 4- and 6-month-old roots, leading to a steeper decline in SRR20 with root age. This result suggests possible increased nutrient uptake efficiency in young fine roots under warmer temperatures. Soil warming significantly increased specific root length (SRL) but not root tissue nitrogen concentration (RTN). The variation in SRR20 between warming treatments, but not across root ages, was predicted by SRL and RTN individually or together. Our findings conclusively indicate that soil warming increased the respiration cost of young fine roots, which was predicted by adjusting for SRL and RTN, indicating that Chinese fir may adopt a faster fine root turnover strategy to enhance nutrient uptake and soil exploitation under warmer temperatures. Future studies should simultaneously investigate age-related root respiration and nutrient uptake in warming experiments to better understand the effects of warming on root metabolic activity., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Due to massive energetic investments in woody support structures, trees are subject to unique physiological, mechanical, and ecological pressures not experienced by herbaceous plants. Despite a wealth of studies exploring trait relationships across the entire plant kingdom, the dominant traits underpinning these unique aspects of tree form and function remain unclear. Here, by considering 18 functional traits, encompassing leaf, seed, bark, wood, crown, and root characteristics, we quantify the multidimensional relationships in tree trait expression. We find that nearly half of trait variation is captured by two axes: one reflecting leaf economics, the other reflecting tree size and competition for light. Yet these orthogonal axes reveal strong environmental convergence, exhibiting correlated responses to temperature, moisture, and elevation. By subsequently exploring multidimensional trait relationships, we show that the full dimensionality of trait space is captured by eight distinct clusters, each reflecting a unique aspect of tree form and function. Collectively, this work identifies a core set of traits needed to quantify global patterns in functional biodiversity, and it contributes to our fundamental understanding of the functioning of forests worldwide., (© 2022. The Author(s).)

Key Message: Among the five cassava isoforms (MeAPL1-MeAPL5), MeAPL3 is responsible for determining storage root starch content. Degree of storage root postharvest physiological deterioration (PPD) is directly correlated with starch content. AGPase is heterotetramer composed of two small and two large subunits each coded by small gene families in higher plants. Studies in cassava (Manihot esculenta) identified and characterized five isoforms of Manihot esculenta ADP-glucose pyrophosphorylase large subunit (MeAPL1-MeAPL5) and employed virus induced gene silencing (VIGS) to show that MeAPL3 is the key isoform responsible for starch and dry matter accumulation in cassava storage roots. Silencing of MeAPL3 in cassava through stable transgenic lines resulted in plants displaying significant reduction in storage root starch and dry matter content (DMC) and induced a distinct phenotype associated with increased petiole/stem angle, resulting in a droopy leaf phenotype. Plants with reduced starch and DMC also displayed significantly reduced or no postharvest physiological deterioration (PPD) compared to controls and lines with high DMC and starch content. This provides strong evidence for direct relationships between starch/dry matter content and its role in PPD and canopy architecture traits in cassava., (© 2020. The Author(s).)

Feedback between plants and soil microbial communities can be a powerful driver of vegetation dynamics. Plants elicit changes in the soil microbiome that either promote or suppress conspecifics at the same location, thereby regulating population density-dependence and species co-existence. Such effects are often attributed to the accumulation of host-specific antagonistic or beneficial microbiota in the rhizosphere. However, the identity and host-specificity of the microbial taxa involved are rarely empirically assessed. Here we review the evidence for host-specificity in plant-associated microbes and propose that specific plant-soil feedbacks can also be driven by generalists. We outline the potential mechanisms by which generalist microbial pathogens, mutualists and decomposers can generate differential effects on plant hosts and synthesize existing evidence to predict these effects as a function of plant investments into defence, microbial mutualists and dispersal. Importantly, the capacity of generalist microbiota to drive plant-soil feedbacks depends not only on the traits of individual plants but also on the phylogenetic and functional diversity of plant communities. Identifying factors that promote specialization or generalism in plant-microbial interactions and thereby modulate the impact of microbiota on plant performance will advance our understanding of the mechanisms underlying plant-soil feedback and the ways it contributes to plant co-existence., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

Root exudation is a key process that determines rhizosphere functions and plant-soil relationships. The present study was conducted with the objectives to (1) determine the root morphology of sessile oak seedlings in relation to drought, (2) assess root exudation and its response to drought, and (3) detect possible changes in the activity of soil enzymes in response to drought enhancement. In the experiment, sessile oak seedlings (Quercus petraea Matt.) were used, and two variants of substrate moisture (25% humidity-dry variant and 55% humidity-fresh variant) on which oaks grew were considered. Exudates were collected using a culture-based cuvette system. Results confirmed the importance of drought in shaping the morphology of roots and root carbon exudation of sessile oak. The oak roots in the dry variant responded with a higher increment in length. In the case of roots growing in higher humidity, a higher specific root area and specific root length were determined. Experimental evidence has demonstrated decreased root exudation under dry conditions, which can lead to a change in enzyme activity. In the study, enzyme activity decreased by 90% for β-D-cellobiosidase (CB), 50% for β-glucosidase (BG) and N-acetyl-β-D-glucosaminidase (NAG), 20% for β-xylosidase (XYL) decreased by, and the activity of arylsulphatase (SP) and phosphatase (PH) decreased by 10%., (© 2022. The Author(s).)

Plants are often exposed to multiple herbivores and densities of these attackers (or corresponding damage intensities) often fluctuate greatly in the field. Plant-mediated interactions vary among herbivore species and with changing feeding intensity, but little is known about how herbivore identity and density interact to determine plant responses and herbivore fitness. Here, we investigated this question using Triadica sebifera (tallow) and two common and abundant specialist insect herbivores, Bikasha collaris (flea beetle) and Heterapoderopsis bicallosicollis (weevil). By manipulating densities of leaf-feeding adults of these two herbivore species, we tested how variations in the intensity of leaf damage caused by flea beetle or weevil adults affected the performance of root-feeding flea beetle larvae and evaluated the potential of induced tallow root traits to predict flea beetle larval performance. We found that weevil adults consistently decreased the survival of flea beetle larvae with increasing leaf damage intensities. In contrast, conspecific flea beetle adults increased their larval survival at low damage then decreased larval survival at high damage, resulting in a unimodal pattern. Chemical analyses showed that increasing leaf damage from weevil adults linearly decreased root carbohydrates and increased root tannin, whereas flea beetle adults had opposite effects as weevil adults at low damage and similar effects as them at high damage. Furthermore, across all feeding treatments, flea beetle larval survival correlated positively with concentrations of carbohydrates and negatively with concentration of tannin, suggesting that root primary and secondary metabolism might underlie the observed effects on flea beetle larvae. Our study demonstrates that herbivore identity and density interact to determine systemic plant responses and plant-mediated effects on herbivores. In particular, effects are species-specific at low densities, but converge at high densities. These findings emphasize the importance of considering herbivore identity and density simultaneously when investigating factors driving plant-mediated interactions between herbivores, which advances our understanding of the structure and composition of herbivore communities and terrestrial food webs., (© 2022 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

One of the characteristic aspects of odour sensing in humans is the activation of olfactory receptors in a slightly different manner in response to different enantiomers. Here, we focused on whether plants showed enantiomer-specific response similar to that in humans. We exposed Arabidopsis seedlings to methanol (control) and (+)- or (-)-borneol, and found that only (+)-borneol reduced the root length. Furthermore, the root-tip width was more increased upon (+)-borneol exposure than upon (-)-borneol exposure. In addition, root-hair formation was observed near the root tip in response to (+)-borneol. Auxin signalling was strongly reduced in the root tip following exposure to (+)-borneol, but was detected following exposure to (-)-borneol and methanol. Similarly, in the root tip, the activity of cyclin B1:1 was detected on exposure to (-)-borneol and methanol, but not on exposure to (+)-borneol, indicating that (+)-borneol inhibits the meristematic activity in the root. These results partially explain the (+)-borneol-specific reduction in the root length of Arabidopsis. Our results indicate the presence of a sensing system specific for (+)-borneol in Arabidopsis.