Inroads into saline-alkaline stress response in plants: unravelling morphological, physiological, biochemical, and molecular mechanisms.

MAIN CONCLUSION: This article discusses the complex network of ion transporters, genes, microRNAs, and transcription factors that regulate crop tolerance to saline-alkaline stress. The framework aids scientists produce stress-tolerant crops for smart agriculture. Salinity and alkalinity are frequently coexisting abiotic limitations that have emerged as archetypal mediators of low yield in many semi-arid and arid regions throughout the world. Saline-alkaline stress, which occurs in an environment with high concentrations of salts and a high pH, negatively impacts plant metabolism to a greater extent than either stress alone. Of late, saline stress has been the focus of the majority of investigations, and saline-alkaline mixed studies are largely lacking. Therefore, a thorough understanding and integration of how plants and crops rewire metabolic pathways to repair damage caused by saline-alkaline stress is of particular interest. This review discusses the multitude of resistance mechanisms that plants develop to cope with saline-alkaline stress, including morphological and physiological adaptations as well as molecular regulation. We examine the role of various ion transporters, transcription factors (TFs), differentially expressed genes (DEGs), microRNAs (miRNAs), or quantitative trait loci (QTLs) activated under saline-alkaline stress in achieving opportunistic modes of growth, development, and survival. The review provides a background for understanding the transport of micronutrients, specifically iron (Fe), in conditions of iron deficiency produced by high pH. Additionally, it discusses the role of calcium in enhancing stress tolerance. The review highlights that to encourage biomolecular architects to reconsider molecular responses as auxiliary for developing tolerant crops and raising crop production, it is essential to (a) close the major gaps in our understanding of saline-alkaline resistance genes, (b) identify and take into account crop-specific responses, and (c) target stress-tolerant genes to specific crops.

Bioaccumulation efficacy and physio-morphological adaptations in response to iron and aluminium contamination of Indian camphorweed (Pluchea indica L.) using different growth substrates.

The aim of this study was to assess the removal capability of Fe/Al contamination of Indian camphorweed (Pluchea indica; hereafter, P. indica) using different growth substrates (100% sand, gardening soil, vermiculite, and zeolite). In addition, the study aimed at observing the physio-morphological adaptation strategies of P. indica under excess Fe/Al levels in a controlled greenhouse environment. After a 4-week treatment, P. indica plants under excess Fe in the 100% sand substrate exhibited signs of decay and eventually death. In contrast, the growth performances of P. indica under gardening soil substrate remained sustained even when exposed to Fe/Al stress. Under zeolite substrate, Fe in the root tissues was 23.1 and 34.7 mg g-1 DW after 1 and 4 weeks of incubation, respectively. In addition, Al in the root tissues also increased to 1.54 mg g-1 DW after 1 week and 1.59 mg g-1 DW after 4 weeks, when subjected to 20 mM Al treatment. Zeolite was observed to be a promising substrate to regulate the uptake of Fe (3.31 mg plant-1) and Al (0.51 mg plant-1) by the root tissues. The restriction of Fe and Al in the root and a low translocation to the leaf organ was indicated by a low translocation factor (< 1.0). High Fe concentrations in the root and leaf tissues negatively affected root elongation, and the net photosynthetic rate decreased by > 40% compared to positive control. Gas exchange parameters and leaf temperature were found the most sensitive to Fe/Al stress. Moreover, the limited transpiration rate under Fe/Al stress caused an increase of the leaf temperature and crop stress index. The findings suggest that P. indica grown using zeolite substrate may serve as a good model system for constructed wetlands, storing excess Al in the root tissues without any significant growth inhibition.

Alleviation of water-deficit stress in turmeric plant (Curcuma longa L.) using phosphate solubilizing rhizo-microbes inoculation.

The objective of this study was to assess the effects of phosphate solubilizing rhizo-microbes inoculants on nutrient balance, physiological adaptation, growth characteristics, and rhizome yield traits as well as curcuminoids yield at the secondary-rhizome initiation stage of turmeric plants, subsequently subjected to water-deficit (WD) stress. Phosphorus contents in the leaf tissues of Talaromyces aff. macrosporus and Burkholderia sp. (Bruk) inoculated plants peaked at 0.33 and 0.29 mg g-1 DW, respectively, under well-watered (WW) conditions; however, phosphorus contents declined when subjected to WD conditions (p ≤ 0.05). Similarly, potassium and calcium contents reached their maximum values at 5.33 and 3.47 mg g-1 DW, respectively, in Burk inoculated plants under WW conditions, which contributed to sustained rhizome fresh weight even when exposed to WD conditions (p ≤ 0.05). There was an increase in free proline content in T. aff. macrosporus and Burk inoculated plants under WD conditions, which played a crucial role in controlling leaf osmotic potential, thereby stabilizing leaf greenness and maximum quantum yield of PSII. As indicators of drought stress, there were noticeable restrictions in stomatal gas exchange parameters, including net photosynthetic rate, stomatal conductance, and transpiration rate, accompanied by an increase in leaf temperature. These changes resulted in reduced total soluble sugar levels. Interestingly, total curcuminoids and curcuminoids yield in Burk inoculated plants under WD conditions were retained, especially in relation to rhizome biomass. Burk inoculation in turmeric plants is recommended as a promising technique as it alleviates water-deficit stress, sustains rhizome biomass, and stabilizes curcuminoids yield. Supplementary Information: The online version contains supplementary material available at 10.1007/s13205-024-03922-x.

Screening cotton genotypes for their drought tolerance ability based on the expression level of dehydration-responsive element-binding protein and proline biosynthesis-related genes and morpho-physio-biochemical responses.

Drought stress adversely affects growth, development, productivity, and fiber quality of cotton (Gossypium hirsutum L). Breeding strategies to enhance drought tolerance require an improved knowledge of plant drought responses necessitating proper identification of drought-tolerant genotypes of crops, including cotton. The objective of this study was to classify the selected cotton genotypes for their drought tolerance ability based on morpho-physio-biochemical traits using Hierarchical Ward's cluster analysis. Five genotypes of cotton (Takfa 3, Takfa 6, Takfa 7, Takfa 84-4, and Takfa 86-5) were selected as plant materials, and were grown under well-watered (WW; 98 ± 2% field capacity) and water-deficit (WD; 50 ± 2% field capacity) conditions for 16 days during the flower initiation stage. Data on morpho-physio-biochemical parameters and gene expression levels for these parameters were collected, and subsequently genotypes were classified either as a drought tolerant or drought susceptible one. Upregulation of GhPRP (proline-rich protein), GhP5CS (Δ1-pyrroline-5-carboxylate synthetase), and GhP5CR (Δ1-pyrroline-5-carboxylate reductase) in relation to free proline enrichment was observed in Takfa 3 genotype under WD condition. An accumulation of free proline, total soluble sugar, and potassium in plants under WD conditions was detected, which played a key role as major osmolytes controlling cellular osmotic potential. Magnesium and calcium concentrations were also enriched in leaves under WD conditions, functioning as essential elements and regulating photosynthetic abilities. Leaf greenness, net photosynthetic rate, stomatal conductance, and transpiration rate were also declined under WD conditions, leading to growth retardation, especially aboveground traits of Takfa 6, Takfa 7, Takfa 84-4, and Takfa 86-5 genotypes. An increase in leaf temperature (1.1 - 4.0 °C) and crop water stress index (CWSI > 0.75) in relation to stomatal closure and reduced transpiration rate was recorded in cotton genotypes under WD conditions compared with WW conditions. Based on the increase of free proline, soluble sugar, leaf temperature, and CWSI, as well as the decrease of aboveground growth traits and physiological attributes, five genotypes were categorized into two cluster groups: drought tolerant (Takfa 3) and drought susceptible (Takfa 6, Takfa 7, Takfa 84-4, and Takfa 86-5). The identified drought-tolerant cotton genotype, namely, Takfa 3, may be grown in areas experiencing drought conditions. It is recommended to further validate the yield traits of Takfa 3 under rainfed field conditions in drought-prone environments.

Exogenous glycine betaine alleviates water-deficit stress in Indian pennywort (Centella asiatica) under greenhouse conditions.

Centella asiatica (Indian pennywort) is a green leafy vegetable containing centelloside' (triterpenoid), a key phytochemical component in traditional medicine. Being a glycophytic species, they exhibit decline in growth performance and yield traits when subjected to water-deficit (WD) conditions. Glycine betaine (GB) is a low molecular-weight organic metabolite that plays a crucial role in abiotic stress conditions in higher plants. The objective of this study was to investigate the potential of GB in alleviating water-deficit stress (in terms of morphological and physiological responses) in two different genotypes of Indian pennywort, "Nakhon Pathom" (NP; high centelloside-yielding genotype) and "Pathum Thani" (PT; low centelloside-yielding genotype). The genotypes of Indian pennywort were propagated by stolon cutting and transplanted into plastic bags containing 2 kg of garden soil. At the flower-initiation stage (30 days after transplantation), uniform plant material was treated exogenously with 0 (control), 25, and 50 mM GB at 100 mL per plant (one-time foliar spray) and then divided into two groups, 1) well watered (WW; irrigated daily with 400 mL fresh water; 98% field capacity) and 2) water deficit (WD; withheld water for 14 days; 72% field capacity). Foliar application of GB (25 mM) significantly improved leaf osmotic potential in NP under WD conditions via osmotic adjustment by free proline and fructose. Differences in leaf temperature (Tleaf) between WD and WW in NP were maximized (+ 1.93 °C) and the gap of Tleaf was reduced in the case of 25-50 mM GB application. Similarly, crop water stress index (CWSI) in NP and PT plants under WD condition was significantly increased by 1.95- and 1.86-fold over the control, respectively; however, it was significantly decreased by exogenous GB application. Increasing Tleaf and CWSI in drought-stressed plants was closely related to stomatal closure, leading to reduced gas exchange parameters, i.e., stomatal conductance (gs), transpiration rate (E), net photosynthetic rate (Pn), and intercellular CO2 concentration (Ci), and consequently decreased plant biomass and total centelloside yield. Overall physiological, morphological, and secondary metabolite traits were enhanced in NP under WD conditions using 25 mM GB exogenous application compared with the control. The study highlights the significance of GB in Indian pennywort production under limited water irrigation (water deficit) with higher vegetable yield and phytochemical stabilization.

Iron (Fe) toxicity, uptake, translocation, and physio-morphological responses in Catharanthus roseus.

Iron (Fe) toxicity in plant species depends on the availability of Fe in the soil, uptake ability by the root system, and translocation rate to other parts of the plant. The aim of this study was to assess Fe uptake by root tissues of Catharanthus roseus, translocation rate to leaf tissues, and the impairment of plant physio-morphological characteristics. Fe uptake by the roots (~ 700 µg g-1 DW) of C. roseus was observed during the early exposure period (1 week), and translocation factor from root to shoot was fluctuated as an independent strategy. A high level of Fe content in the root tissues significantly inhibited root length and root dry weight. Under acidic pH condition, an enrichment of Fe in the shoots (~ 400 µg g-1 DW) led to increase in leaf temperature (> 2.5 °C compared to control) and crop stress index (> 0.6), resulting in stomatal closure, subsequently decreasing CO2 assimilation rate and H2O transpiration rate. An increment of CSI in Fe-stressed plants was negatively related to stomatal conductance, indicating stomatal closure with an increase in Fe in the leaf tissues. High Fe levels in the leaf tissues directly induced toxic symptoms including leaf bronzing, leaf spotting, leaf necrosis, leaf chlorosis, and leaf senescence in C. roseus plants. In summary, C. roseus was identified as a good candidate plant for Fe phytoextraction, depending on Fe bioaccumulation, therefore 50 mM Fe treatment was designated as an excess Fe to cause the growth inhibition, especially in the prolonged Fe incubation periods. Supplementary Information: The online version contains supplementary material available at 10.1007/s12298-023-01379-5.

Investigating high throughput phenotyping based morpho-physiological and biochemical adaptations of indian pennywort (Centella asiatica L. urban) in response to different irrigation regimes.

Indian pennywort (Centella asiatica L. Urban; Apiaceae) is a herbaceous plant used as traditional medicine in several regions worldwide. An adequate supply of fresh water in accordance with crop requirements is an important tool for maintaining the productivity and quality of medicinal plants. The objective of this study was to find a suitable irrigation schedule for improving the morphological and physiological characteristics, and crop productivity of Indian pennywort using high-throughput phenotyping. Four treatments were considered based on irrigation schedules (100, 75, 50, and 25% of field capacity denoted by I100 [control], I75, I50, and I25, respectively). The number of leaves, plant perimeter, plant volume, and shoot dry weight were sustained in I75 irrigated plants, whereas adverse effects on plant growth parameters were observed when plants were subjected to I25 irrigation for 21 days. Leaf temperature (Tleaf) was also retained in I75 irrigated plants, when compared with control. An increase of 2.0 °C temperature was detected in the Tleaf of plants under I25 irrigation treatment when compared with control. The increase in Tleaf was attributed to a decreased transpiration rate (R2 = 0.93), leading to an elevated crop water stress index. Green reflectance and leaf greenness remained unchanged in plants under I75 irrigation, while significantly decreased under I50 and I25 irrigation. These decreases were attributed to declined leaf osmotic potential, increased non-photochemical quenching, and inhibition of net photosynthetic rate (Pn). The asiatic acid and total centellosides in the leaf tissues, and centellosides yield of plants under I75 irrigation were retained when compared with control, while these parameters were regulated to maximal when exposed to I50 irrigation. Based on the results, I75 irrigation treatment was identified as the optimum irrigation schedule for Indian pennywort in terms of sustained biomass and a stable total centellosides. However, further validation in the field trials at multiple locations and involving different crop rotations is recommended to confirm these findings.

Toxicity, physiological, and morphological alterations of Indian camphorweed (Pluchea indica) in response to excess copper.

Indian camphorweed (Pluchea indica (L.) Less.) is used as herbal tea due to the presence of volatile aromatic oils and several phytochemical compounds. The aim of this study was to assess the impact of copper (Cu) contamination on the physiology and morphology of P. indica, and the health risks associated with its consumption as tea. The cuttings of P. indica were subjected to 0 mM (control), 5 mM (low Cu), and 20 mM (excess Cu) of CuSO4 treatments for 1, 2, and 4 weeks. Thereafter, Cu contamination as well as physiological and morphological parameters were assessed. Cu accumulation was higher in the root tissues of plants (25.8 folds higher as compared to the leaves) grown under 20 mM CuSO4 for 4 weeks. This increased Cu accumulation resulted in the inhibition of root length, root fresh weight, and root dry weight. Cu concentration was found maximum (1.36 µg g-1 DW) in the leaf tissues under 20 mM Cu exposure for 4 weeks, with the highest target hazard quotient (THQ = 1.85), whereas Cu was not detected in control. Under exposure to 20 mM Cu treatment for 4 weeks, leaf greenness, maximum quantum yield of photosystem II, and photon yield of photosystem II diminished by 21.4%, 16.1%, and 22.4%, respectively, as compared to the control. Leaf temperature was increased by 2.5 °C, and the crop stress index (CSI) exceeded 0.6 when exposed to 20 mM Cu treatment for 2 and 4 weeks; however, the control had a CSI below 0.5. This led to a reduced transpiration rate and stomatal conductance. In addition, the net photosynthetic rate was also found sensitive to Cu treatment, which resulted in decreased shoot and root growth. Based on the key results, it can be suggested that P. indica herbal tea derived from the foliage of plants grown under a 5 mM Cu level (0.75 µg g-1 DW) with a target hazard quotient below one aligns with the recommended dietary intake of Cu in leafy vegetables. The study recommends choosing cuttings from plants with a small canopy as plant material in the greenhouse microclimates to validate the growth performance in the Cu-contaminated soil and simulate the natural shrub architecture and life cycle.

Integrated strength of osmotic potential and phosphorus to achieve grain yield of rice under water deficit by arbuscular mycorrhiza fungi.

Arbuscular mycorrhizal ecosystem provides sustainability to plant integrity under drought situations. However, host plants that survive in drought frequently lose yield. The potential of Funneliformis mosseae (F), Claroideoglomus etunicatum (C), and Acaulospora fovaeta (A) was assessed to evaluate in indica rice cv. Leum Pua during booting stage under 21-day water withholding. The effects of three inoculation types; (i) F, (ii) F + C (FC), and (iii) F + C + A (FCA), on physiological, biochemical, and yield traits were investigated. The three types showed an induced total chlorophyll content in the host as compared to uninoculated plants. Total soluble sugars and free proline were less regulated by FC and FCA inoculated plants than by F inoculated plants under water deficit conditions. However, the FC and FCA inoculations increased phosphorus content, particularly in the shoots of water-stressed plants. In the three inoculations, the FCA dramatically improved plant osmotic potential adaptability under water deficit stress. Furthermore, even when exposed to the water deficit condition, panicle weight, grain number, and grain maturity were maintained in FCA inoculated plants. According to the findings, the increased osmotic potential and phosphorus content of the FCA-inoculated rice plant provide a protection sign against drought stress and will benefit food security in the future.

Aluminum uptake, translocation, physiological changes, and overall growth inhibition in rice genotypes (Oryza sativa) at vegetative stage.

Aluminum (Al) contamination in acidic soil is a major problem in paddy field, causing grain yield loss, especially in central plains of Thailand. The objective of this study was to assess Al content in the root tissues, its translocation to the leaves, and Al toxicity in three genotypes of rice, RD35 (local acidic-tolerant), Azucena (positive-check Al-tolerant), and IR64 (high yielding) under 0 (control) or 1 mM AlCl3 (Al toxicity) at pH 4.5. Al content in the root tissues of rice cv. RD35 under 1 mM AlCl3 was peaked at 4.18 mg g‒1 DW and significantly translocated to leaf tissues (0.35 mg g‒1 DW), leading to reduced leaf greenness (SPAD) (by 44.9% over the control) and declined net photosynthetic rate (Pn) (by 54.5% over the control). In contrast, Al level in cvs. Azucena and IR64 was restricted in the roots (2.12 mg g‒1 DW) with low amount of translocation in the leaf tissues (0.26 mg g‒1 DW), resulting in maintained values of SPAD and Pn. In cv. RD35, root and shoot traits including root length, root fresh weight, shoot height, shoot fresh weight, and shoot dry weight in 1 mM Al treatment were significantly dropped by > 35% over the control, whereas these parameters in cvs. Azucena and IR64 were retained. Based on the results, RD35 rice genotype was identified as Al sensitive as it demonstrated Al toxicity in both aboveground and belowground parts, whereas Azucena and IR64 were found tolerant to 1 mM Al as they demonstrated storage of Al in the root tissues to reduce toxicity in the leaf tissues. The study suggests that root traits, shoot attributes, chlorophyll degradation, and photosynthetic reduction can be successfully employed for the screening of Al-tolerant genotypes in rice breeding programs.

Expression levels of nitrogen assimilation-related genes, physiological responses, and morphological adaptations of three indica rice (Oryza sativa L. ssp. indica) genotypes subjected to nitrogen starvation conditions.

Nitrogen (N) is an essential nutrient available to the plants in form of nitrate and ammonium. It is a macronutrient important for the plant growth and development, especially in cereal crops, which consume it for the production of amino acids, proteins/enzymes, nucleic acids, cell wall complexes, plant hormones, and vitamins. In rice production, 17 kg N uptake is required to produce 1 ton of rice. Considering this, many techniques have been developed to evaluate leaf greenness or SPAD value for assessing the amount of N application in the rice cultivar to maximize the grain yield. The aim of the present study was to investigate the morpho-physiological characteristics and relative expression level of N assimilation in three different rice genotypes (MT2, RD31, KDML105) under 1.00 × (full N), 0.50 × , 0.25 × (N depletion), and 0.00 × (N deficiency) at seedling stage and the morpho-physiological traits and the grain yield attributes under 1.00 × (full N) and 0.25 × (N depletion) were compared. Leaf chlorosis and growth inhibition in rice seedlings under N deficiency were evidently observed. Shoot height, number of leaves, shoot fresh weight, shoot dry weight, and root fresh weight in KDML105 under N deficiency were decreased by 27.65%, 42.11%, 65.44%, 47.90%, and 54.09% over the control (full N). Likewise, leaf greenness was lowest in KDML105 under N deficiency (78.57% reduction over the full N), leading to low photosynthetic abilities. In addition, expression of nitrogen assimilation-related genes, OsNR1, OsGln1;1, and OsGln2, in KDML105 under N depletion were increased within 3 h and then declined after the long incubation period, whereas those were unchanged in cvs. MT2 and RD31. Similarly, relative expression level of OsNADH-GOGAT, OsFd-GOGAT, and OsAspAt1 in KDML105 was peaked when subjected to 0.50 × N for 6 h and then declined after the long incubation period. Moreover, overall growth characters and physiological changes in cv. RD31 at vegetative stage under 0.25 × N were retained better than those in cvs. KDML105 and MT2, resulting in high yield at the harvesting process. In summary, N assimilated-related genes in rice seedlings under N depletion were rapidly regulated within 3-6 h, especially cv. KDML105 and MT2, then downregulated, resulting in physiological changes, growth inhibition, and yield reduction.

Effectiveness of vegetation indices and UAV-multispectral imageries in assessing the response of hybrid maize (Zea mays L.) to water deficit stress under field environment.

Unmanned aerial vehicles (UAVs) equipped with multi-sensors are one of the most innovative technologies for measuring plant health and predicting final yield in field conditions, especially in the water deficit situation in rain-deprived regions. The objective of this investigation was to evaluate the individual plant and canopy-level measurements using UAV imageries in three different genotypes, Suwan4452 (drought-tolerant), Pac339, and S7328 (drought-sensitive) of maize (Zea mays L.) at vegetative and reproductive stages under WW (well-watered) and WD (water deficit) conditions. At the vegetative stage, only CWSI (crop water stress index) of Pac339 and S7328 under WD increased significantly by 1.86- and 1.69-fold over WW, whereas the vegetation indices (EVI2 (Enhanced Vegetation Index 2), OSAVI (Optimized Soil-Adjusted Vegetation Index), GNDVI (Green Normalized Difference Vegetation Index), NDRE (Normalized Difference Red Edge Index), and NDVI (Normalized Difference Vegetation Index)) derived from UAV multi-sensors did not vary. At the reproductive stage, CWSI in drought-sensitive genotype (S7328) under WD increased by 1.92-fold over WW. All the vegetation indices (EVI2, OSAVI, GNDVI, NDRE, and NDVI) of Pac339 and S7328 under WD decreased when compared with those of Suwan4452. NDVI derived from GreenSeeker® handheld and NDVI from UAV data was closely related (R2 = 0.5924). An increase in leaf temperature (Tleaf) and reduction in NDVI of WD stressed maize plants was observed (R2 = 0.5829) leading to yield loss (R2 = 0.5198). In summary, a close correlation was observed between the physiological data of individual plants and vegetation indices of canopy level (collected using a UAV platform) in drought-sensitive genotypes of maize crops under WD conditions, thus indicating its effectiveness in the classification of drought-tolerant genotypes.

Arbuscular mycorrhizal fungi inoculation and phosphorus application improve growth, physiological traits, and grain yield of rice under alternate wetting and drying irrigation.

Climate change and agricultural malpractices are exacerbating drought in many parts of the world causing a substantial agricultural production loss. The improvement of drought tolerance in rice is crucial for maintaining productivity and ensuring global food security. Alternate wetting and drying (AWD) irrigation along with plant-microbe interaction through arbuscular mycorrhizal fungi (AMF) is a potential approach for enhancing rice production through AMF-induced up-regulation of tolerance and resilience against drought stress. Therefore, the ameliorative role of AMF inoculation and phosphorus (P) application on growth, physiological traits, and grain yield of rice was evaluated under water stress imposed through AWD irrigation. A factorial experiment consisting of four fertilizer treatments where the P percentage varied along with the recommended dose of nitrogen (N) with or without AMF inoculation (P100 as the control, P100 + AMF, P75 + AMF, and P50 + AMF), three soil water potential levels (0, -15, and -30 kPa), and two cultivation methods (wet direct seeding and transplanting) was conducted in a polyhouse. The subscript values of 100, 75, and 50 under P represent 100%, 75%, and 50% of the recommended field application dose. Data were collected on selected growth parameters, physiological traits, levels of mycorrhizal colonization, yield and its components, and water productivity of rice. The results revealed that P100 + AMF inoculated plants had 11%, 14%, 74%, and 54% higher leaf greenness, leaf relative water content, net photosynthetic rate, and grain yield, respectively, for wet direct-seeded plants at reduced soil water potential (-30 kPa) compared with non-inoculated plants (P100). Free proline accumulation gradually enhanced with decreasing soil water potential, and it was maximized by 77% at -30 kPa compared with 0 kPa for P50 + AMF (for transplanted plants). Free proline accumulation was also higher with decreasing soil water potential in AMF-inoculated plants than non-inoculated plants regardless of cultivation methods. Leaf osmotic potential was reduced by -0.5 to -1.2 MPa at -30 kPa compared with 0 kPa under different fertilizer doses. However, AMF inoculation (P100 + AMF and P75 + AMF) improved leaf osmotic potential of plants under severe water stress (-30 kPa) maintained through AWD irrigation resulting in better osmotic adjustment than non-inoculated plants. AMF inoculation improved the response of most of the evaluated physiological traits of rice and enhanced grain yield with higher P availability (even with a 25% reduction in its recommended dose) in the rhizosphere under drought stress. Thus, it can be concluded that AMF inoculation coupled with judicious P management is a promising approach for improving physiological and biochemical traits, grain yield, and water productivity of rice under AWD irrigation regardless of cultivation methods.

Determination of traits responding to iron toxicity stress at different stages and genome-wide association analysis for iron toxicity tolerance in rice (Oryza sativa L.).

Rice is the staple food for more than half of the world's population. Iron toxicity limits rice production in several regions of the world. Breeding Fe-tolerant rice varieties is an excellent approach to address the problem of Fe toxicity. Rice responds differently to Fe toxicity at different stages. Most QTLs associated with Fe toxicity have been identified at the seedling stage, and there are very few studies on Fe toxicity across different stages. In this study, we investigated agro-morphological and physiological traits in response to Fe toxicity in a rice diversity panel at seedling, vegetative, and reproductive stages and applied GWAS to identify QTLs/genes associated with these traits. Among agro-morphological and physiological parameters, leaf bronzing score (LBS) is a key parameter for determining Fe toxicity response at all stages, and SDW could be a promising parameter at the seedling stage. A total of 29 QTLs were identified on ten chromosomes. Among them, three colocalized QTLs were identified on chromosome 5, 6, and 11. Several QTLs identified in this study overlapped with previously identified QTLs from bi-parental QTL mapping and association mapping. Two genes previously reported to be associated with iron homeostasis were identified, i.e., LOC_Os01g72370 (OsIRO2, OsbHLH056) and LOC_Os04g38570 (OsABCB14). In addition, based on gene-based haplotype analysis, LOC_Os05g16670 was identified as a candidate gene for the colocalized QTL on chromosome 5 and LOC_Os11g18320 was identified as a candidate gene for the colocalized QTL on chromosome 11. The QTLs and candidate genes identified in this study could be useful for rice breeding programs for Fe toxicity tolerance.

Matching of Nitrogen Enhancement and Photosynthetic Efficiency by Arbuscular Mycorrhiza in Maize (Zea mays L.) in Relation to Organic Fertilizer Type.

In the present study, Funneliformis mosseae (FM), Claroideoglomus etunicatum (CE), and Acaulospora foveata (AF) were inoculated to hybrid maize (Zea mays L. cv. CP888®). Upregulation of nitrogen levels were dependent on the type of mycorrhiza (AMF). Photosynthetic efficiency (Fv/Fm) and water content in FM- and AF-inoculated plants were elevated, resulting in promotion of leaf area and shoot biomass. N content in the shoot and root tissues of the FM-inoculated plants increased by 21% and 30% over the control. A positive correlation between biochemical, physiological, and morphological parameters using Pearson's coefficient was demonstrated. A decline in lipid peroxidation was noticed in the FM-inoculated plants. In addition, we investigated the potential of N fertilizer application in combination with FM inoculation in maize plants. The FM-inoculated plants with organic O_LT, a chicken manure fertilizer, increased N content in the host shoots by 73% over the control, leading to improved Fv/Fm as a physiological adaptation strategy. The FM and the O_LT on the regulation of the N enhancement and photosynthetic efficiency of the hybrid maize should further be validated in field trials in different environments for sustainability.

Evaluation of curcuminoids, physiological adaptation, and growth of Curcuma longa under water deficit and controlled temperature.

Turmeric (Curcuma longa L.; Zingiberaceae), an economically important crop and a major spice in Indian cuisine, produces natural yellow color (curcumin) as well as curcuminoids which are widely utilized in traditional and modern medicinal practices. During the turmeric culture, the fluctuations of precipitation and seasonal changes in the whole life cycle play a major role, especially water shortage and decreasing temperature (in winter season), leading to rhizome dormancy under extreme weather conditions. The objective of this investigation was to understand how the water deficit and reduced temperature affect turmeric growth, physiological adaptation, quantity, and quality of turmeric rhizomes. Four-month-old turmeric plants were subjected to four treatments, namely normal temperature and well-watered (RT-WW), or water-deficit (RT-WD) conditions in the greenhouse, 25 °C controlled temperature and well-watered (CT-WW), or water-deficit (CT-WD) conditions in glasshouse. Leaf osmotic potential considerably declined in 30 days CT-WD treatment, leading to chlorophyll degradation by 26.04%, diminution of maximum quantum yield of PSII (Fv/Fm) by 23.50%, photon yield of PSII (ΦPSII) by 29.01%, and reduction of net photosynthetic rate (Pn) by 89.39% over CT-WW (control). After 30 days water withholding, fresh- and dry-weights of rhizomes of turmeric plants grown under CT-WD declined by 30-50% when compared with RT-WW conditions. Subsequently, curcuminoid content was reduced by 40% over RT-WW plants (control), whereas transcriptional expression levels of curcuminoids-related genes (CURS1, CURS2, CURS3, and DCS) were upregulated in CT-WD conditions. In summary, the water withholding and controlled temperature (constant at 25 °C day/night) negatively affected turmeric plants as abiotic stresses tend to limit overall plant growth performances and curcuminoid yield.

Expression levels of genes involved in metal homeostasis, physiological adaptation, and growth characteristics of rice (Oryza sativa L.) genotypes under Fe and/or Al toxicity.

Acid sulphate soil contains high amounts of iron (Fe) and aluminum (Al), and their contamination has been reported as major problems, especially in rainfed and irrigated lowland paddy fields. Rice is sensitive to Fe and Al grown in acid soil (pH < 5.5), leading to growth inhibition and grain yield loss. The objective of this study was to evaluate Fe and/or Al uptake, translocation, physiological adaptation, metal toxicity, and growth inhibition in rice genotypes grown in acid soil. Fe and Al in the root tissues of all rice genotypes were enriched depending on the exogenous application of either Fe or Al in the soil solution, leading to root growth inhibition, especially in the KDML105 genotype. Expression level of OsYSL1 in KDML105 was increased in relation to metal uptake into root tissues, whereas OsVIT2 was downregulated, leading to Fe (50.3 mg g-1 DW or 13.1 folds over the control) and Al (4.8 mg g-1 DW or 2.2 folds over the control) translocation to leaf tissues. Consequently, leaf greenness (SPAD), net photosynthetic rate (Pn), stomatal conductance (gs), and transpiration rate (E) in the leaf tissues of genotype KDML105 under Fe + Al toxicity significantly declined by 28.4%, 35.3%, 55.6%, and 51.6% over the control, respectively. In Azucena (AZU; Fe/Al tolerant), there was a rapid uptake of Fe and Al by OsYSL1 expression in the root tissues, but a limited secretion into vacuole organelles by OsVIT2, leading to a maintenance of low level of toxicity driven by an enhanced accumulation of glutathione together with downregulation of OsGR expression level. In addition, Fe and Al restrictions in the root tissues of genotype RD35 were evident; therefore, crop stress index (CSI) of Fe + Al-treated plants was the maximum, leading to an inhibition of gs (53.6% over the control) and E (49.0% over the control). Consequently, free proline, total phenolic compounds, and ascorbic acid in the leaf tissues of rice under Fe + Al toxicity significantly increased by 3.2, 1.2, and 1.5 folds over the control, respectively, indicating their functions in non-enzymatic antioxidant defense. Moreover, physiological parameters including leaf temperature (Tleaf) increment, high level of CSI (>0.6), SPAD reduction, photon yield of PSII (ΦPSII) diminution, Pn, gs, and E inhibition in rice genotype IR64 (Fe/Al-sensitive) under Fe + Al treatment were clearly demonstrated as good indicators of metal-induced toxicity. Our results on Fe- and/or Al-tolerant screening to find out the candidate genotypes will contribute to present screening and breeding efforts, which in turn help increase rice production in the Fe/Al-contaminated acid soil under lowland conditions.

Physio-morphological traits and osmoregulation strategies of hybrid maize (Zea mays) at the seedling stage in response to water-deficit stress.

Drought has been identified as a major factor restricting maize productivity worldwide, especially in the rainfed areas. The objective of the present study was to investigate the physiological adaptation strategies and sugar-related gene expression levels in three maize (Zea mays L.) genotypes with different drought tolerance abilities (Suwan4452, drought tolerant as a positive check; S7328, drought susceptible as a negative check; Pac339, drought susceptible) at the seedling stage. Ten-day old seedlings of maize genotypes were subjected to (i) well-watered (WW) or control and (ii) water-deficit (WD) conditions. Leaf osmotic potential of cv. S7328 under WD was significantly decreased by 1.35-1.45 folds compared with cv. Pac339 under WW, whereas it was retained in cv. Suwan4452, which utilized total soluble sugars as the major osmolytes for maintaining leaf greenness, Fv/Fm, ΦPSII, and stomatal function (Pn, net photosynthetic rate; gs, stomatal conductance; and E, transpiration rate). Interestingly, sucrose degradation (65% over the control) in cv. Pac339 under WD was evident in relation to the downregulation of the ZmSPS1 level, whereas glucose enrichment (1.65 folds over the control) was observed in relation to the upregulation of ZmSPS1 and ZmSUS1. Moreover, CWSI (crop water stress index), calculated from leaf temperature of stressed plants, was negatively correlated with E, gs, and Pn. Overall, growth characteristics, aboveground and belowground parts, in the drought-susceptible cv. Pac339 and cv. S7328, were significantly decreased (> 25% over the control), whereas these parameters in the drought-tolerant cv. Suwan4452 were unaffected. The study validates the use of leaf temperature, CWSI, Pn, gs, and E as sensitive parameters and overall growth characters as effective indices for drought tolerance screening in maize genotypes at the seedling stage. However, further experiments are required to validate the results observed in this study under field conditions.

In vitro acclimatization of Curcuma longa under controlled iso-osmotic conditions.

In vitro acclimatization has been validated as the successful key to harden the plantlets before transplanting to ex vitro conditions. In the present study, we investigated the potential of different sugar types (glucose, fructose, galactose, sucrose) in regulating morphological, physiological and biochemical strategies, survival percentage and growth performance, and rhizome traits of turmeric under iso-osmotic potential. Leaf greenness (SPAD value) in acclimatized plantlets (4% glucose; -1.355 MPa osmotic potential) of 'ST018' was retained and greater than in 'PB009' by 1.69-fold, leading to maintain high Fv/Fm (maximum quantum yield of PSII), ΦPSII (photon yield of PSII) and Pn (net photosynthetic rate) levels, and retained shoot height, leaf length, leaf width, shoot fresh weight and shoot dry weight after one month upon transplanting to ex vitro conditions. In addition, Pn, Ci (intracellular CO2), gs (stomatal conductance) and E (transpiration rate) in acclimatized plantlets (6% sucrose; -1.355 MPa osmotic potential) of 'PB009' were stabilized as physiological adapted strategies, regulating the shoot and root growth and fresh and dry weights of mini-rhizome. Interestingly, the accumulation of total curcuminoids in mini-rhizome derived from 6% sucrose acclimatized plantlets of 'ST018' was greater than in 'PB009' by 3.76-fold. The study concludes that in vitro acclimation of turmeric 'PB009' and 'ST018' using 6% sucrose and 4% glucose, respectively, promoted percent survival, physiological adaptations, and overall growth performances under greenhouse conditions.

Characterization of macrophytes for Na+ removal in synthetic Na-salt solution batch under greenhouse conditions.

Sodium salt contamination in the fresh water due to industrial effluents, underground rock salts and inland aquaculture is a major concern needs to be remediated, and subsequently recycled as sustainable bioeconomic strategy. Treatment of saline wastewater requires efficient, cost-effective, rapid, and green technologies, so as to mitigate the negative impacts of salinity on agricultural land. Green technology of phytodesalination is proposed to reduce salinity in the wastewater using salt tolerant plant species. present study was designed with an aim to investigate the sodium (Na+) removal capacity of salt tolerant and high biomass producing macrophytes on synthetic saline wastewater. Sesuvium portulacastrum (sea purslane), Pluchea indica (Indian camphorweed), Typha angustifolia (narrow leaf cattail) and Heliconia psittacorum (heliconia) were collected, cultivated in the greenhouse, subsequently treated with 0 (control) and 217 mM NaCl (salt stress) for 4 weeks. Overall growth performance, physiological change and Na+ removal rate in root and leaf tissues of the candidate plant species were measured. Plants were able to maintain their growth and physiological abilities except for shoot height in T. angustifolia (reduced by 13.7%) and chlorophyll content in S. portulacastrum (reduced by 64%). Major accumulation of Na+ was recorded in the shoots of S. portulacastrum and P. indica (halophytic plant species) and the roots of T. angustifolia and H. psittacorum (glycophytic plant species). Since T. angustifolia and H. psittacorum have high plant biomass, they showed higher Na+ removal efficiency at 4.4% and 5.7%, respectively; whereas due to lower plant biomass, S. portulacastrum and P. indica resulted in the removal of only 0.6 and 0.8% Na+ from the batch, respectively. Based on the information from this investigation, the selected candidate plant species can further be studied in the constructed wetland together with the controlled environments including optimized flowrate, vertical or horizontal flow system, plant densities and Na-removal rate in relation to swamp habitat.Novelty statement: T. angustifolia and H. psittacorum have high plant biomass, they showed higher Na+ removal efficiency at 4.4% and 5.7%, respectively; whereas due to lower plant biomass, S. portulacastrum and P. indica resulted in removal of only 0.6 and 0.8% Na+ from the batch. Based on the information from this investigation, the selected candidate plant species can further be studied in the constructed wetland together with the controlled environments including optimized flowrate, vertical or horizontal flow system, plant densities and Na-removal rate in relation to swamp habitat.