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Single-cell C4 (SCC4) plants with bienertioid anatomy carry out photosynthesis in a single cell. Chloroplast movement is the underlying phenomenon, where chloroplast unusual positioning 1 (CHUP1) plays a key role. This study aimed to characterize CHUP1 and CHUP1-like proteins in an SCC4 photosynthetic plant, Bienertia sinuspersici. Also, a comparative analysis of SCC4 CHUP1 was made with C3, C4, and CAM model plants including an extant basal angiosperm, Amborella. The CHUP1 gene exists as a single copy from the basal angiosperms to SCC4 plants. Our analysis identified that Chenopodium quinoa, a recently duplicated allotetraploid, has two copies of CHUP1. In addition, the numbers of CHUP1-like and its associated proteins such as CHUP1-like_a, CHUP1-like_b, HPR, TPR, and ABP varied between the species. Hidden Markov Model analysis showed that the gene size of CHUP1-like_a and CHUP1-like_b of SCC4 species, Bienertia, and Suaeda were enlarged than other plants. Also, we identified that CHUP1-like_a and CHUP1-like_b are absent in Arabidopsis and Amborella, respectively. Motif analysis identified several conserved and variable motifs based on the orders (monocot and dicot) as well as photosynthetic pathways. For instance, CAM plants such as pineapple and cactus shared certain motifs of CHUP1-like_a irrespective of their distant phylogenetic relationship. The free ratio model showed that CHUP1 maintained purifying selection, whereas CHUP1-like_a and CHUP1-like_b have adaptive functions between SCC4 plants and quinoa. Similarly, rice and maize branches displayed functional diversification on CHUP1-like_b. Relative gene expression data showed that during the subcellular compartmentalization process of Bienertia, CHUP1 and actin-binding proteins (ABP) genes showed a similar pattern of expression. Altogether, the results of this study provide insight into the evolutionary and functional details of CHUP1 and its associated proteins in the development of the SCC4 system in comparison with other C3, C4, and CAM model plants.

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Plants deposit lignin in the secondary cell wall as a common response to drought and pathogen attacks. Cell wall localised multicopper oxidase family enzymes LACCASES (LACs) catalyse the formation of monolignol radicals and facilitate lignin formation. We show an upregulation of the expression of several LAC genes and a downregulation of microRNA397 (CamiR397) in response to natural drought in chickpea roots. CamiR397 was found to target LAC4 and LAC17L out of twenty annotated LACs in chickpea. CamiR397 and its target genes are expressed in the root. Overexpression of CamiR397 reduced expression of LAC4 and LAC17L and lignin deposition in chickpea root xylem causing reduction in xylem wall thickness. Downregulation of CamiR397 activity by expressing a short tandem target mimic (STTM397) construct increased root lignin deposition in chickpea. CamiR397-overexpressing and STTM397 chickpea lines showed sensitivity and tolerance, respectively, towards natural drought. Infection with a fungal pathogen Macrophomina phaseolina, responsible for dry root rot (DRR) disease in chickpea, induced local lignin deposition and LAC gene expression. CamiR397-overexpressing and STTM397 chickpea lines showed more sensitivity and tolerance, respectively, to DRR. Our results demonstrated the regulatory role of CamiR397 in root lignification during drought and DRR in an agriculturally important crop chickpea.

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Salt stress is an ever-increasing stressor that affects both plants and humans. Therefore, developing strategies to limit the undesirable effects of salt stress is essential. Sodium ion exclusion is well known for its efficient salt-tolerance mechanism. The High-affinity K+ Transporter (HKT) excludes excess Na+ from the transpiration stream. This study identified and characterized the HKT protein family in Bienertia sinuspersici, a single-cell C4 plant. The HKT and Salt Overly Sensitive 1 (SOS1) expression levels were examined in B. sinuspersici and Arabidopsis thaliana leaves under four different salt stress conditions: 0, 100, 200, and 300 mM NaCl. Furthermore, BsHKT1;2 was cloned, thereby producing stable transgenic Brassica rapa. Our results showed that, compared to A. thaliana as a glycophyte, the HKT family is expanded in B. sinuspersici as a halophyte with three paralogs. The phylogenetic analysis revealed three paralogs belonging to the HKT subfamily I. Out of three copies, the expression of BsHKT1;2 was higher in Bienertia under control and salt stress conditions than in A. thaliana. Stable transgenic plants overexpressing 35S::BsHKT1;2 showed higher salt tolerance than non-transgenic plants. Higher biomass and longer roots were observed in the transgenic plants under salt stress than in non-transgenic plants. This study demonstrates the evolutionary and functional differences in HKT proteins between glycophytes and halophytes and associates the role of BsHKT1;2 in imparting salt tolerance and productivity.

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Abiotic stressors such as drought and heat predispose chickpea plants to pathogens of key importance leading to significant crop loss under field conditions. In this study, we have investigated the influence of drought and high temperature on the incidence and severity of dry root rot disease (caused by Macrophomina phaseolina) in chickpea, under extensive on- and off-season field trials and greenhouse conditions. We explored the association between drought tolerance and dry root rot resistance in two chickpea genotypes, ICC 4958 and JG 62, with contrasting resistance to dry root rot. In addition, we extensively analyzed various patho-morphological and root architecture traits altered by combined stresses under field and greenhouse conditions in these genotypes. We further observed the role of edaphic factors in dry root rot incidence under field conditions. Altogether, our results suggest a strong negative correlation between the plant water relations and dry root rot severity in chickpeas, indicating an association between drought tolerance and dry root rot resistance. Additionally, the significant role of heat stress in altering the dynamics of dry root rot and the importance of combinatorial screening of chickpea germplasm for dry root rot resistance, drought, and heat stress have been revealed.

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Drought plays a central role in increasing the incidence and severity of dry root rot (DRR) disease in chickpea. This is an economically devastating disease, compromising chickpea yields particularly severely in recent years due to erratic rainfall patterns. Macrophomina phaseolina (formerly Rhizoctonia bataticola) is the causal agent of DRR disease in the chickpea plant. The infection pattern in chickpea roots under well-watered conditions and drought stress are poorly understood at present. This study provides detailed disease symptomatology and the characteristics of DRR fungus at morphological and molecular levels. Using microscopy techniques, the infection pattern of DRR fungus in susceptible chickpea roots was investigated under well-watered and drought-stress conditions. Our observations suggested that drought stress intensifies the progression of already ongoing infection by weakening the endodermal barrier and overall defense. Transcriptomic analysis suggested that the plant's innate immune defense program is downregulated in infected roots when subjected to drought stress. Furthermore, genes involved in hormonal regulation are differentially expressed under drought stress. These findings provide hints in terms of potential chickpea genes to target in crop improvement programs to develop climate-change-resilient cultivars.[Formula: see text] Copyright © 2022 The Author(s). This is an open access article distributed under the CC BY-NC-ND 4.0 International license.

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Chickpea is an essential crop for protein nutrition and is grown around the world in rain-fed conditions. However, chickpea cultivation is under threat due to emerging diseases favored by drought stress. Dry root rot (DRR), an economically devastating disease, is an example. Chickpea-specific strains of a necrotic fungal phytopathogen, Macrophomina phaseolina, cause DRR. Microsclerotia of this fungus, which are capable of withstanding harsh environmental conditions, serve as primary inoculum. Initial symptoms are scattered necrotic spots in roots, progressing to rotting and withering lateral roots, accompanied by prematurely dried, straw-colored foliage. The recent rise in global temperature and worsening of drought spells have aggravated DRR outbreaks in chickpea. To date, DRR epidemiology has not been clarified in detail. Also, the literature lacks clarity on M. phaseolina taxonomy, morphology, disease progression, and diagnosis. In this article, research progress on patterns of DRR occurrence in the field and belowground and aboveground symptoms are clarified. In addition, the current understanding of taxonomy and management practices is elaborated. We also summarize knowledge of the impact of drought and high temperature on DRR severity. Furthermore, we provide future perspectives on the importance of host resistance, quantitative trait loci identification, and genotype screening for the identification of resistant genotypes. The article proposes new research priorities and a corresponding plan for the mitigation of DRR.

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PREMISE: A comprehensive field-based screening protocol is lacking for dry root rot (DRR) disease in chickpea, which is caused by Macrophomina phaseolina (formerly referred to as Rhizoctonia bataticola). Here, we describe a protocol for establishing a sick plot for DRR to enable disease assessment of a large number of chickpea plants during the natural growing season. METHODS AND RESULTS: We used a chickpea plot with >30% DRR incidence, and enriched the inoculum by cultivating highly susceptible chickpea plant genotypes and incorporating infected plant material into the soil. The chickpea plants were then subjected to infection in developed sick plots with various levels of soil moisture under natural field conditions. CONCLUSIONS: Our protocol provides a robust way to impose M. phaseolina infection on chickpea plants under natural field conditions and to investigate plant responses to the infection at morphological, physiological, and molecular levels. This method can also be used to screen for other soil-borne diseases in a variety of plants.

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Rhizoctonia bataticola causes dry root rot (DRR), a devastating disease in chickpea (Cicer arietinum). DRR incidence increases under water deficit stress and high temperature. However, the roles of other edaphic and environmental factors remain unclear. Here, we performed an artificial neural network (ANN)-based prediction of DRR incidence considering DRR incidence data from previous reports and weather factors. ANN-based prediction using the backpropagation algorithm showed that the combination of total rainfall from November to January of the chickpea-growing season and average maximum temperature of the months October and November is crucial in determining DRR occurrence in chickpea fields. The prediction accuracy of DRR incidence was 84.6% with the validation dataset. Field trials at seven different locations in India with combination of low soil moisture and pathogen stress treatments confirmed the impact of low soil moisture on DRR incidence under different agroclimatic zones and helped in determining the correlation of soil factors with DRR incidence. Soil phosphorus, potassium, organic carbon, and clay content were positively correlated with DRR incidence, while soil silt content was negatively correlated. Our results establish the role of edaphic and other weather factors in chickpea DRR disease incidence. Our ANN-based model will allow the location-specific prediction of DRR incidence, enabling efficient decision-making in chickpea cultivation to minimize yield loss.

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Dry root rot (DRR) disease is an emerging biotic stress threat to chickpea cultivation around the world. It is caused by a soil-borne fungal pathogen, Rhizoctonia bataticola. In the literature, comprehensive and detailed step-by-step protocols on disease assays are sparse. This article provides complete details on the steps involved in setting up a blotting paper technique for quickly screening genotypes for resistance to DRR. The blotting paper technique is easy and less expensive. Another method, based on the sick pot approach, is a mimic of natural infection and can be applied to study the interacting components-plant, pathogen, and environment-involved in the disease triangle. Moreover, in nature, DRR occurs mostly in rainfed chickpea cultivation areas, where soil moisture recedes as crop growth advances. Drought stress is known to predispose chickpea plants to DRR disease. Pathomorphological and molecular understanding of plant-pathogen interaction under drought stress can pave the way for the identification of elite DRR-resistant varieties from the chickpea germplasm pool. This article provides a stepwise methodology for the preparation of a sick pot and subsequent disease assay. Overall, the information presented herein will help researchers prepare R. bataticola fungal inoculum, maintain this pathogen, set up the blotting paper technique, prepare sick culture and sick pot, and assess pathogen infection in chickpea plants.

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Drought stress and pathogen infection simultaneously occur in the field. In this study, the interaction of these two stresses with chickpea, their individual and combined effect and the net impact on plant growth and yield traits were systematically assessed under field and confined pot experiments. The field experiments were conducted for four consecutive years from 2014-15 to 2017-18 at different locations of India. Different irrigation regimes were maintained to impose mild to severe drought stress, and natural incidence of the pathogen was considered as pathogen stress. We observed an increased incidence of fungal diseases namely, dry root rot (DRR) caused by Rhizoctonia bataticola, black root rot (BRR) caused by Fusarium solani under severe drought stress compared to well-irrigated field condition. Similar to field experiments, pot experiments also showed severe disease symptoms of DRR and BRR in the presence of drought compared to pathogen only stress. Overall, the results from this study not only showed the impact of combined drought and DRR stress but also provided systematic data, first of its kind, for the use of researchers.

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Global warming leads to the concurrence of a number of abiotic and biotic stresses, thus affecting agricultural productivity. Occurrence of abiotic stresses can alter plant-pest interactions by enhancing host plant susceptibility to pathogenic organisms, insects, and by reducing competitive ability with weeds. On the contrary, some pests may alter plant response to abiotic stress factors. Therefore, systematic studies are pivotal to understand the effect of concurrent abiotic and biotic stress conditions on crop productivity. However, to date, a collective database on the occurrence of various stress combinations in agriculturally prominent areas is not available. This review attempts to assemble published information on this topic, with a particular focus on the impact of combined drought and pathogen stresses on crop productivity. In doing so, this review highlights some agriculturally important morpho-physiological traits that can be utilized to identify genotypes with combined stress tolerance. In addition, this review outlines potential role of recent genomic tools in deciphering combined stress tolerance in plants. This review will, therefore, be helpful for agronomists and field pathologists in assessing the impact of the interactions between drought and plant-pathogens on crop performance. Further, the review will be helpful for physiologists and molecular biologists to design agronomically relevant strategies for the development of broad spectrum stress tolerant crops.