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Curvularia lunata (No. CLST-01), a fungal pathogen isolated from the threeleaf arrowhead (Sagittaria trifolia L.), has been proposed as a potential mycoherbicide for grass weeds. This paper investigated the physiological and biochemical effects of CLST-01 phytotoxic ethyl acetate fungi extract on the leaves of the threeleaf arrowhead. The results showed that the ethyl acetate fungi extract from CLST-01 can accelerate damage to the cell membrane, increase the production of malondialdehyde, and damage the cellular structure, which could decrease the number of chloroplasts after 96 h treatments. In addition, the content of chlorophyll was reduced by 49.5%, and the net photosynthetic rate, stomatal conductance, and transpiration rate were inhibited. The rates of inhibition were 90.13%, 83.74%, and 79.31%, respectively, and the intercellular CO2 concentration increased by 51.87% on Day 9 after treatment with a concentration of 200 µg/mL. In summary, the phytotoxic ethyl acetate fungal extract from C. lunata CLST-01 can inhibit the photosynthesis of the threeleaf arrowhead leaves, destroy the ultrastructure of leaves, and affect the growth of this invasive weed. Therefore, it has the potential to be developed into a mycoherbicide for weed control in crops as a natural photosynthetic inhibitor.

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Sagittaria trifolia control is threatened by the emergence of resistance to acetolactate synthase (ALS)-inhibiting herbicides. Hence, we systematically uncovered the molecular mechanism of resistance to the main herbicide (bensulfuron-methyl) in Liaoning Province from target-site and non-target-site resistance perspectives. The suspected resistant population (TR-1) exhibited high-level resistance. A new amino acid substitution (Pro-197-Ala) in resistant Sagittaria trifolia for ALS was detected, and the molecular docking results showed that the spatial structure of ALS changed significantly after the substitution, manifested by an increase in the number of contacted amino acid residues and the disappearance of hydrogen bonds. Dose-response test of transgenic Arabidopsis thaliana further demonstrated that the Pro-197-Ala substitution conferred bensulfuron-methyl resistance. The assays found that the sensitivity of the ALS enzyme in TR-1 to this herbicide was decreased in vitro; and this population had developed resistance to other types of ALS-inhibiting herbicides. Furthermore, the resistance of TR-1 to bensulfuron-methyl was significantly alleviated after co-treatment with a P450-inhibitor (malathion). TR-1 metabolized bensulfuron-methyl significantly faster than sensitive population (TS-1) did, but this gap was narrowed after malathion treatment. Overall, the resistance of Sagittaria trifolia to bensulfuron-methyl was derived from the mutation of the target-site gene and the enhancement of the P450s-mediated detoxification metabolism.

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Background: Sagittaria trifolia Linn. is a widespread macrophyte in Asia and southeast Europe and cultivated in parts of Asia. Although a few genomic studies have been conducted for S. trifolia var. sinensis, a crop breed, there is limited genomic information on the wild species of S. trifolia. Effective microsatellite markers are also lacking. Objective: To assemble transcriptome sequence and develop effective EST-SSR markers for S. trifolia. Methods: Here we developed microsatellite markers based on tri-, tetra-, penta-, and hexa-nucleotide repeat sequences by comparatively screening multiple transcriptome sequences of eleven individuals from ten natural populations of S. trifolia. Results: A total of 107,022 unigenes were de novo assembled, with a mean length of 730 bp and an N50 length of 1,378 bp. The main repeat types were mononucleotide, trinucleotide, and dinucleotide, accounting for 55.83%, 23.51%, and 17.56% of the total repeats, respectively. A total of 86 microsatellite loci were identified with repeats of tri-, tetra-, penta-, and hexa-nucleotide. For SSR verification, 28 polymorphic loci from 41 randomly picked markers were found to produce stable and polymorphic bands, with the number of alleles per locus ranging from 2 to 11 and a mean of 5.2. The range of polymorphic information content (PIC) of each SSR locus varied from 0.25 to 0.80, with an average of 0.58. The expected heterozygosity ranged from 0.29 to 0.82, whereas the observed heterozygosity ranged from 0.25 to 0.90. Conclusion: The assembled transcriptome and annotated unigenes of S. trifolia provide a basis for future studies on gene functions, pathways, and molecular mechanisms associated with this species and other related. The newly developed EST-SSR markers could be effective in examining population genetic structure, differentiation, and parentage analyses in ecological and evolutionary studies of S. trifolia.

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Weeds tend to develop resistance to herbicides with time. Understanding the resistance mechanisms evolved by weeds would help manage weed infestation. Sagittaria trifolia, a paddy weed found in the rice fields of Liaoning, China, has developed resistance to bensulfuron-methyl, causing severe yield losses in rice. This study deciphers the underlying mechanisms in terms of non-target-site resistance toward bensulfuron-methyl. We compared the ability of glutathione S-transferase (GST) mediated detoxification metabolism and reactive oxygen species (ROS) scavenging between sensitive (NHS) and resistant (NHR) populations of S. trifolia. The resistance ratio of NHR was 210; but the ratio was significantly decreased after GST-inhibitor treatment (44.9). This indicated that a GST-mediated enhancement of detoxification metabolism stimulated the development of resistance. Similarly, higher GST activity was observed in NHR; but the activity equaled that of NHS after GST-inhibitor treatment. However, treatment with the GST-inhibitor did not completely reverse bensulfuron-methyl resistance in NHR, indicating that additional factors contributed to herbicide resistance in these plants. We observed a rapid increase in H2O2 and malondialdehyde accumulation in the case of NHS after bensulfuron-methyl application, whereas those of NHR remained relatively stable, implying that NHR exhibited higher ROS-scavenging capacity under herbicide stress. Further, NHR showed higher glutathione and ascorbic acid contents and higher activities of glutathione reductase and dehydrogenase reductase, all of which contribute towards herbicide resistance in these plants. Our results indicate that GST-mediated detoxification metabolism of bensulfuron-methyl and ROS scavenging capacity contributed to the development of resistance in S. trifolia.

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Sagittaria trifolia tuber is an aquatic vegetable. In this work, microwave-assisted enzymatic extraction (MEE) was used to extract S. trifolia tuber polysaccharides (STTPs). Optimum conditions were complex enzyme of 2 %, liquid-to-solid ratio of 43 : 1 mL g-1 , microwave power of 506 W, and time of 8 min, under which STTPs yield was 36.22±0.69 %, higher than those of other methods. STTPs were sulfated polysaccharides with sulfur valence of S6+ . STTPs comprised mannose, glucose, galactose, and arabinose at a mole ratio of 3.69 : 19.33 : 6.21 : 1.00, molecular weights of 3606 kDa and 149.6 kDa, particle size of 220 nm, and zeta potential of -5.02 mV. The surface of STTPs was full of bumps and holes, and abundant in O1s and non-functionalized C1s. STTPs would scavenge reactive oxygen species with advantage. It would provide an efficient MEE method to obtain antioxidant STTPs, also a clue for extracting polysaccharides from starch-rich crops.

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Acetohydroxy acid synthase (AHAS)-inhibiting herbicides are one of the most commonly used herbicides for controlling the growth of Sagittaria trifolia L. in paddy fields in Northeastern China. In this study, we collected five suspected resistant populations of S. trifolia (R1-R5) from three different provinces of Northeastern China. The results of whole-plant bioassays revealed that those populations showed high level of resistance to bensulfuron-methyl with resistance index (GR50 R/S) ranging from 39.90 to 88.50. The results of AHAS-activity assays were consistent with the results of the whole-plant bioassays. The AHAS gene analysis showed that R2 and R3 populations contained Pro-197-Leu mutations that were highly resistant to penoxsulam; R1 and R4 populations contained Pro-197-Ser mutations that were highly resistant to bispyribac­sodium; R5 population contained Trp-574-Leu mutation that showed high resistance to IMI, PT, PTB and SU herbicides. The AHAS with resistance mutations showed less sensitivity to feedback inhibition by BCAAs and R genotypes had increased free BCAAs.

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Sagittaria trifolia is an aquatic plant that is distributed worldwide. The edible tuber part of S. trifolia is a very common and popular vegetable in China. The aim of the present review is to discuss the discovery of nutraceuticals from S. trifolia tuber by reviewing its major constituents, food processing, food products, and health-promoting benefits. Sagittaria trifolia tuber comprises a series of nutritional and bioactive constituents, including dietary fibers, amino acids, minerals, starches, non-starch polysaccharides, diterpenoids, colchicine, phenols, and organic acids. Food processing affects its flavor, biocomponents, and bioactivity. Numerous S. trifolia tuber-based food products and nutraceuticals have been developed, but new categories of products and the anticipated functions still need to be explored. The non-starch polysaccharides could be the central ingredients that contribute to the plant's antioxidant, hepatoprotective, hypoglycemic, lipid-regulating, and immunostimulatory properties. Of these, antioxidant and hepatoprotective effects have been thoroughly investigated. Procedures for the extraction and purification of polysaccharides influence their health-promoting actions. Overall, S. trifolia tuber is an underutilized aquatic vegetable species that is an emerging subject for nutraceutical research. © 2020 Society of Chemical Industry.

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Sagittaria trifolia is a medicinal foodstuff of China and East Asia belonging to the family Alismataceae. Samples of S. trifolia tubers were collected from Meihekow, Siping, Jilin, Harbin and Wuchang from Northeast China. The current study was aimed to evaluate the qualitative and quantitative analysis, antioxidant activity, biochemical analysis and chemical composition of different populations of S. trifolia. By using Folin-Ciocalteu, aluminium chloride colourimetric and 1,1-diphenyl-1-picrylhydrazyl (DPPH), total phenol and flavonoids content and antioxidant activity was analysed. Furthermore, chemical composition, biochemical analysis and mineral substances were also determined. The results showed the presence of flavonoids, phenols, saponins, tannins, glycosides and steroids except for alkaloids and terpenoids by qualitative analysis. Quantitative analysis revealed that highest total phenol, flavonoids content and antioxidant potential identified from Meihekow, i.e., 2.307 mg GAE/g, 12.263 mg QE/g and 77.373%, respectively. Gas chromatography-mass spectrometry results showed the presence of 40 chemical compounds corresponding to 99.44% of total extract that might be responsible for antioxidant properties. Mineral and biochemical analysis revealed the presence of calcium, magnesium, potassium, sodium, iron, copper, zinc and, carbohydrate, protein, fibre and fat contents, respectively. Interestingly, all S. trifolia populations collected from different locations possess similar composition. The dietary values, phytoconstituents, antioxidant activities and nutritional and curative chemical compounds of S. trifolia are beneficial for the nutritherapy of human beings.

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Aquatic macrophytes are considered to be promising in controlling harmful cyanobacterial blooms. In this research, an aqueous extract of Sagittaria trifolia tubers was prepared to study its inhibitory effect on Microcystis aeruginosa in the laboratory. Several physiological indices of M. aeruginosa, in response to the environmental stress, were analyzed. Results showed that S. trifolia tuber aqueous extract significantly inhibited the growth of M. aeruginosa in a concentration-dependent way. The highest inhibition rate reached 90% after 6 day treatment. The Chlorophyll-a concentration of M. aeruginosa cells decreased from 343.1 to 314.2µg/L in the treatment group. The activities of superoxide dismutase and peroxidase and the content of reduced glutathione in M. aeruginosa cells initially increased as a response to the oxidative stress posed by S. trifolia tuber aqueous extract, but then decreased as time prolonged. The lipid peroxidation damage of the cyanobacterial cell membranes was reflected by the malondialdehyde level, which was notably higher in the treatment group compared with the controls. It was concluded that the oxidative damage of M. aeruginosa induced by S. trifolia tuber aqueous extract might be one of the mechanisms for the inhibitory effects.

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Arrowhead (Sagittaria trifolia var. sinensis) is a source of starch worldwide, but arrowhead starch has been rarely studied. In this work, starch was separated from arrowhead corm. The morphology and physicochemical properties of starch were then investigated and compared among 3 different arrowhead varieties (Purple-corm, Hongta, and Japanese). Results showed that starches from the 3 varieties similarly featured an oval shape containing a visible polarization cross, a CA -type crystalline structure, and an ordered structure in the external granule region. However, starch content, granule size, crystal characteristics, and pasting properties differed among the 3 varieties. Japanese arrowhead exhibited the highest starch content and degree of ordered structure in the external granule region, as well as onset, peak, and final gelatinization temperature. Purple-corm arrowhead starch demonstrated the highest amylose content and relative degree of crystallinity, smallest granule size, and lowest swelling power and solubility. Purple-corm arrowhead starch also showed the highest gelatinization enthalpy, as well as peak, trough, final, and setback viscosities. This starch further presented the lowest breakdown viscosity and degree of hydrolysis by HCl and porcine pancreatic α-amylase. These findings can provide useful references for arrowhead variety selection in food and nonfood industries.

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Sagittaria trifolia L. is one of the most serious weeds in paddy fields in northeast of China and cannot be controlled effectively by bensulfuron-methyl in recent years. In this study, two suspected resistant S. trifolia populations (R1 and R2) were collected in Liaoning province of China. Whole-plant dose-response studies showed that R1 and R2 were highly resistant to bensulfuron-methyl, with the GR50 R/S ratios of 76.99 and 49.94 respectively. In vitro acetolactate synthase (ALS) assays revealed that resistance was due to reduced sensitivity of the ALS to bensulfuron-methyl inhibition, with I50 R/S ratios of 81.86 and 67.48 for R1 and R2, respectively. Total ALS activity was similar for the S and R2 populations, whereas the R1 population displayed significantly higher ALS activity than did the S population. The mutations Pro-197-Leu and Pro-197-Ser were identified in the ALS gene of the R1 and R2 populations, respectively. This is the first report examining bensulfuron-resistant S. trifolia in Liaoning province, China. The Pro197 mutation is likely responsible for resistance to bensulfuron-methyl in S. trifolia populations.

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The phytoremediation potential and responses of Sagittaria trifolia to diesel were investigated. In order to elucidate the biochemical and physiological responses of S. trifolia to diesel, the chlorophyll content, root vitality, soluble protein content and antioxidant enzymes activity (peroxidase (POD), catalase (CAT) and antioxidant enzymes superoxide dismutase (SOD)) were determined in the plant tissues after 50 d of diesel treatment. The results showed the presence of S. trifolia significantly improved the removal ratios of diesel, from 21∼36% in the control soils to 54∼85% in the planted soils. The chlorophyll content, root vitality and soluble protein content all increased at low diesel concentration, then decreased at high diesel concentration. The activities of CAT and POD exhibited peak values at 5 g·kg(-1) diesel treatment and declined at higher diesel concentrations. However, the activity of SOD kept stable at lower diesel concentration (1 and 5 g·kg(-1)), and also declined at higher diesel concentration. Collectively, S. trifolia had the ability to tolerate certain amount of diesel, but when the concentration was up to 10 g·kg(-1), the growth of S. trifolia would be restrained. The results also showed that variation of antioxidant enzyme activity was an important response in plants to diesel pollution.

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• Premise of the study: Carpel closure is commonly considered as a key innovation in angiosperms; however, some families continue to exhibit a variety of forms of incomplete carpel closure. The carpel of Sagittaria species contains an unusual structure. In this study, we confirm the closure type of the carpel of Sagittaria trifolia and discuss its development and evolution.• Methods: Scanning electron microscopy and light microscopy of semithin sections were used to observe the development and the mature structure of the carpel. Pollen tube growth in the carpel and seed germination in the achene was also studied.• Key results: During late carpel development, the middle parts of the carpel margins underwent postgenital fusion. However, at maturity the lowest and uppermost parts of the carpel margins remained open. The mature carpel was incompletely closed and contained a secretion-filled canal, which extended from the stigma to the opening at the carpel base. At that site, pollen tubes could either grow to the ovule or exit the carpel and grow to other carpels. The basal opening also served as an exit point for the seedling to emerge.• Conclusions: Incomplete carpel closure by S. trifolia differs from the closure types recorded in previous studies because two entrances link the ovary in the carpel to the outside environment. This type of carpel closure occurs as a result of the lack of fusion of the carpel margins at the base of the carpel and could improve the seed set and seedling germination of S. trifolia.