Melatonin and dopamine alleviate waterlogging stress in apples by recruiting beneficial endophytes to enhance physiological resilience

Melatonin and dopamine can potentially prevent waterlogging stress in apples.The current study investigated the mechanism by which melatonin and dopamine alleviate apple waterlogging stress.This study demonstrated that melatonin and dopamine alleviated waterlogging by removing reactive oxygen species(ROS),and that the nitric oxide(NO)content and nitrate reductase(NR)activity were significantly correlated.Melatonin and dopamine were also found to recruit different candidate beneficial endophytes(melatonin:Novosphingobium,Propionivibrio,and Cellvibrio;dopamine:Hydrogenophaga,Simplicispira,Methyloversatilis,Candidatus_Kaiserbacteria,and Humicola),and these endophytes were significantly and positively correlated with plant growth.Network analyses showed that melatonin and dopamine significantly affected the endophytic bacterial and fungal communities under waterlogging stress.The metabolomic results showed that melatonin and dopamine led to waterlogging resistance by upregulating the abundance of beneficial substances such as amino acids,flavonoids,coumarins,and organic acids.In addition,melatonin and dopamine regulated the physicochemical properties of the soil,which altered the endophyte community and affected plant growth.The co-occurrence network demonstrated close and complex relationships among endophytes,metabolites,soil,and the plants.Our results demonstrate that melatonin and dopamine alleviate waterlogging stress in apples by recruiting beneficial endophytes to enhance physiological resilience.This study provides new insights into how melatonin and dopamine alleviate stress and a theoretical basis for synergistic beneficial microbial resistance to waterlogging stress.

SiRAP2-12,a Positive Regulatory Factor,Effectively Improves the Waterlogging Tolerance of Foxtail Millet(Setaria italica)

Foxtail millet(Setaria italica)growth was inhibited because of waterlogging stress,which has caused yield reduc-tion.ERF family plays an important role to plant adversity tolerance.In our study,we obtained 19,819 differential expressed genes(DEGs)between the two treatments based on the RNA-seq sequencing of foxtail millet of water-logging stress.Furthermore,a total of 28 ERF family members were obtained,which have a complete open read-ing frame.We studied the evolution and function of SiERF family and how they affected the waterlogging tolerance.It was found that SiERF1A/B/C(GenBank ID:OR775217,OR775219,OR775218)and SiRAP2-12(GenBank ID:OR775216)have similar functions to the known waterlogging tolerance genes of other plants.Among them,the SiRAP2-12 expression was obviously significantly up-regulated in foxtail millet after 5d water-logging stress.After SiRAP2-12 was silenced,the activity of defense enzymes in millet decreased significantly.In details,superoxide dismutase(SOD),catalase(CAT)and peroxidase(POD),the osmotic regulator proline(Pro),and the activity of the anaerobic respiratory enzyme alcohol dehydrogenase(ADH)content were decreased by 78.61%,29.52%,79.95%,19.41%and 54.77%,respectively.In contrast,the relative electrical conductivity contents(REC),malondialdehyde(MDA),and hydrogen peroxide(H_(2)O_(2))of the foxtail millet subjected to virus-induced gene silencing clearly increased by 1.03-fold,36.09%,and 15.21%,respectively.The content of sodium(Na^(+))in the SiRAP2-12-silenced foxtail millet also increased,but that of potassium(K^(+))decreased.Interestingly,we found that ethylene content was significantly reduced.Further,the SiAOC1 expression,an essential gene for ethylene synthesis,was inhibited in SiRAP2-12-silenced foxtail millet after waterlogging stress.Taken together,we hypothesized that SiRAP2-12 might be a positive regulator of millet tolerance to waterlogging stress.

Responses of photosynthetic characteristics and leaf senescence in summer maize to simultaneous stresses of waterlogging and shading

A field experiment was performed to investigate the physiological mechanism of the simultaneous stresses of waterlogging and shading on leaf photosynthetic and senescence during three growth stages of summer maize.The responses of leaf gas exchange parameters and antioxidant enzyme activities of the summer maize hybrids Denghai 605(DH605)to waterlogging(W),shading(S),and their combination(W+S)for 6 days at the third leaf stage(V3),the sixth leaf stage(V6),and the tasseling stage(VT)were recorded.Shading,waterlogging,and their combination disturbed the activities of protective enzymes and increased the contents of H2O2and O-2,accelerating leaf senescence and disordering photosynthetic characteristics.Under waterlogging,shading and their combination,leaf Pn,the photo-assimilates and grain yield was decreased.The greatest reduction for waterlogging and the combined stresses occurred at V3 and that for shading stress occurred at VT.The individual and combined stresses reduced the activities of protective enzymes and inhibited photosynthesis,reducing the accumulation of photosynthetic compounds and thereby yield.Waterlogging and the combined stresses at the V3 stage showed the greatest effect on leaf photosynthetic and senescence,followed by the V6 and VT stages.The greatest effect for shading stress occurred at VT,followed by the V6 and V3 stages,and the combined influence of shading and waterlogging was greater than that of either single stress.

Effect of Fatigue Loading on the Mechanical Properties and Resistance of High-strength Straight-hole Recycled Pervious Concrete to Rainstorm-based Waterlogging

A novel high-strength straight-hole recycled pervious concrete(HSRPC)for the secondary highway pavement was prepared in this paper.This study aimed to investigate the effect of porosity(0.126%,0.502%,and 1.13%),vehicle loading stress level(0.5 and 0.8)and service life on the resistance to rainstormbased waterlogging of HSRPC under fatigue loading.The mechanical properties of HSRPC in terms of flexural strength and dynamic elastic modulus were studied.The waterlogging resistance of HSRPC was described by surface water depth and drainage time.The microstructure of HSRPC were observed with scanning electron microscopy(SEM).Results showed that although the dynamic elastic modulus and flexural strength of HSRPC decreased with the increasing number of fatigue loading,the flexural strength of HSRPC was still greater than5 MPa after design service life of 20 years.After 2.5×10^(5)times of fatigue loading,the permeability coefficient of HSRPC with a porosity of 0.502%and 1.13%increased by 18.4%and 22.9%,respectively;while the permeability coefficient of HSRPC with 0.126%porosity dropped to 0.35 mm/s.The maximum surface water depth of HSRPC with a porosity of 0.126%,0.502%,and 1.13%were 8,5 and 4 mm,respectively.SEM results showed that fatigue loading expanded the number and width of cracks around the tiny pores in HSRPC.

Pore Characteristic Design Method of High-strength Pervious Concrete Based on the Mechanical Properties and Rainstorm Waterlogging Resistance

High-strength pervious concrete(HSPC) with porosity ranging from 0.08% to 2.011% was prepared. The mechanical properties and rainstorm waterlogging resistance of HSPC were evaluated,and a design method of HSPC pore characteristics(porosity and pore diameter) based on the mechanical properties and rainstorm waterlogging resistance was proposed. The results showed that the reduction of effective cross-sectional area caused by artificial channels was the main factor affecting flexural strength but had limited influence on compressive strength. Compared with the concrete matrix without artificial channels,the compressive strength of HSPC with porosity of 2.011% decreased by 7.4%, while the flexural strength decreased by 48.3%. The permeability coefficient of HSPC can reach 16.35 mm/s even at low porosity(2.011%).HSPC can meet the requirements of no rainstorm waterlogging, even if exposed to 100-year rainstorms. When the mechanical properties and rainstorm waterlogging resistance are compromised, the recommended porosity ranges from 1.1% to 3.5%, and the recommended pore diameter ranges from 0.8 to 2.7 mm.

Identification of stable quantitative trait loci underlying waterlogging tolerance post-anthesis in common wheat(Triticum aestivum)

Waterlogging is a growing threat to wheat production in high-rainfall areas.In this study,a doubled haploid(DH) population developed from a cross between Yangmai 16(waterlogging-tolerant) and Zhongmai895(waterlogging-sensitive) was used to map quantitative trait loci(QTL) for waterlogging tolerance using a high-density 660K single-nucleotide polymorphism(SNP) array.Two experimental designs,waterlogging concrete tank(CT) and waterlogging plastic tank(PT),were used to simulate waterlogging during anthesis in five environments across three growing seasons.Waterlogging significantly decreased thousand-kernel weight(TKW) relative to non-waterlogged controls,although the degree varied across lines.Three QTL for waterlogging tolerance were identified on chromosomes 4AL,5AS,and 7DL in at least two environments.All favorable alleles were contributed by the waterlogging-tolerant parent Yangmai16.QWTC.caas-4AL exhibited pleiotropic effects on both enhancing waterlogging tolerance and decreasing plant height.Six high-confidence genes were annotated within the QTL interval.The combined effects of QWTC.caas-4AL and QWTC.caas-5AS greatly improved waterlogging tolerance,while the combined effects of all three identified QTL(QWTC.caas-4AL,QWTC.caas-5AS,and QWTC.caas-7DL) exhibited the most significant effect on waterlogging tolerance.Breeder-friendly kompetitive allele-specific PCR(KASP) markers(K_AX_111523809,K_AX_108971224,and K_AX_110553316) flanking the interval of QWTC.caas-4AL,QWTC.caas-5AS,and QWTC.caas-7DL were produced.These markers were tested in a collection of 240 wheat accessions,and three superior polymorphisms of the markers distributed over 67elite cultivars in the test population,from the Chinese provinces of Jiangsu,Anhui,and Hubei.The three KASP markers could be used for marker-assisted selection(MAS) to improve waterlogging tolerance in wheat.

Enhancing waterlogging tolerance in cotton through agronomic practices

Recent publications have highlighted significant progress in utilizing agronomic interventions to alleviate waterlogging stress in cotton production.Based on these advancements,we provide a concise comment on the effects and underlying mechanisms of various strategies such as utilizing stress-tolerant cotton varieties,applying nitric oxide(NO),and implementing ridge intertillage.Finally,we recommend a combination of measures to enhance cotton’s ability to withstand waterlogging and reduce yield losses.

Changes of Physiological Characters of Wheat After Water-logging at Booting and Relations Between Physiological Characters and Waterlogging Tolerance

Effects of waterlogging on some physiological characters and relationship between waterlogging tolerance and some characters of their own were studied by using 12 varieties of wheat. Results showed waterlogging made the content of malondialdehyde in flag leaf increase, made the root vigor, the content of chlorophyll, the net photosynthesis rate, the nitrate reductase activity, the relative water content of flag leaf decrease, the content of organic matter change obviously, and then the yield per plant declined. The correlations

Preliminary Study on the Waterlogging Tolerance of 116 Corn Materials

Maize is one of the most important food crops in the world.With the global warming,waterlogging stress has become an important abiotic stress factor that affects crop growth,including maize.Waterlogging seriously affects 10%of the arable land and can lead to a 15%-80%reduction in crop yield[1].In this study,115 inbred line materials commonly used in spring maize planting areas in the Jianghan Plain,Hubei,and maize inbred line B73 with complete genome information,were collected and stressed by waterlogging for two weeks in the seven-leaf and one-heart stage,and the survival rate was statistically compared and analyzed,aiming to screen germplasms with strong waterlogging tolerance for the genetic improvement of waterlogging tolerance of Hubei maize lines.

Changes of Antioxidative Enzymes and Lipid Peroxidation in Leaves and Roots of Waterlogging-Tolerant and Waterlogging-Sensitive Maize Genotypes at Seedling Stage

To better understand the physiological and biochemical mechanisms of waterlogging tolerance, waterlogging effects on lipid peroxidation and the activity of antioxidative enzymes were investigated in leaves and roots of two maize genotypes, HZ32 （waterlogging-tolerant） and K12 （waterlogging-sensitive）. Potted maize plants were waterlogged at the second leaf stage under glasshouse conditions. Leaves and roots were harvested 1 d before and 2, 4, 6, 8 and 10 d after the start of waterlogging treatment. Through comparing the activities of superoxide dismutase （SOD）, ascorbate peroxidase （APX）, glutathione reductase （GR）, catalase （CAT） and guaiacol peroxidase （POD） between waterlogging-tolerant and waterloggingsensitive genotype, we deduced that CAT was the most important H2O2 scavenging enzyme in leaves, while APX seemed to play a key role in roots. POD, APX, GR and CAT activities in conjunction with SOD seem to play an essential protective role in the O2^- and H2O2 scavenging process. Lipid peroxidation was enhanced significantly only in K12 （P 〈 0.001） and there was no difference （P 〉 0.05） in HZ32 up to 6 d after waterlogging stress. These results indicated that oxidative stress may play an important role in waterlogging-stressed maize plants and that the greater protection of HZ32 leaves and roots from waterlogging-induced oxidative damage results, at least in part, through the maintenance of increased antioxidant enzyme activity.

QTLs for Waterlogging Tolerance at Germination and Seedling Stages in Population of Recombinant Inbred Lines Derived from a Cross Between Synthetic and Cultivated Wheat Genotypes

Waterlogging is a widespread limiting factor for wheat production throughout the world. To identify quantitative trait loci （QTLs） associated with waterlogging tolerance at early stages of growth, survival rate （SR）, germination rate index （GRI）, leaf chlorophyll content index （CCI）, root length index （RLI）, plant height index （PHI）, root dry weight index （RDWI）, shoot dry weight index （SDWI）, and total dry weight index （DWI） were assessed using the International Triticeae Mapping Initiative （ITMI） population W7984/Opata85. Significant and positive correlations were detected for all traits in this population except RLI. A total of 32 QTLs were associated with waterlogging tolerance on all chromosomes except 3A, 3D, 4B, 5A, 5D, 6A, and 6D. Some of the QTLs explained large proportions of the phenotypic variance. One of these is the QTL for GRI on 7A, which explained 23.92% of the phenotypic variation. Of them, 22 alleles from the synthetic hexaploid wheat W7984 contributed positively. These results suggested that synthetic hexaploid wheat W7984 is an important genetic resource for waterlogging tolerance in wheat. These alleles conferring waterlogging tolerance at early stages of growth in wheat could be utilized in wheat breeding for improving waterlogging tolerance.

Comparison of transcriptomes undergoing waterlogging at the seedling stage between tolerant and sensitive varieties of Brassica napus L.

RNA sequencing of the sensitive GH01 variety of Brassica napus L. seedling roots under 12 h of waterlogging was compared with previously published data of the ZS9 tolerant variety to unravel genetic mechanisms of waterlogging tolerance beyond natural variation. A total of 2 977 genes with similar expression patterns and 17 genes with opposite expression patterns were identiifed in the transcription proifles of ZS9 and GH01. An additional 1 438 genes in ZS9 and 1 861 genes in GH01 showed strain speciifc regulation. Analysis of the overlapped genes between ZS9 and GH01 revealed that waterlogging tolerance is determined by ability to regulate genes with similar expression patterns. Moreover, differences in both gene expression proifles and abscisic acid (ABA) contents between the two varieties suggest that ABA may play some role in waterlogging tolerance. This study identiifes a subset of candidate genes for further functional analysis.

Screening Methods for Waterlogging Tolerance at Maize (Zea mays L.) Seedling Stage

Waterlogging strongly affects agronomic performance of maize （Zea mays L.）. In order to investigate the suitable selection criteria of waterflooding tolerant genotypes, and identify the most susceptible stage and the best continuous treatment time to waterlogging, 20 common maize inbred lines were subjected to successive artificial waterflooding at seedling stage, and waterlogging tolerance coefficient （WTC） was used to screen waterflooding tolerant genotypes. In addition, peroxidase （POD） activities and malondialdehyde （MDA） contents were measured for 6 of 20 lines. The results showed that the second leaf stage （V2） was the most susceptible stage, and 6 d after waterflooding was the best continuous treatment time. Dry weight （DW） of both shoots and roots of all lines were significantly reduced at 6 d time-point of waterlogging, compared to control. POD activities and MDA contents were negatively and significantly correlated, and the correlation coefficient was -0.9686 （P 〈 0.0001）. According to the results, WTC of shoot DW can be used for practical screening as a suitable index, which is significantly different from control and waterlogged plants happened 6 d earlier. Furthermore, leaf chlorosis, MDA content and POD activities could also be used as reference index for material screening. The implications of the results for waterlogging-tolerant material screening and waterlogging-tolerant breeding have been discussed in maize.

Combining Ability and Genetic Effects of Germination Traits of Brassica napus L.Under Waterlogging Stress Condition

Cross combinations from six rapeseed cultivars and lines were evaluated under waterlogging stress condition in order to understand the genetic mechanism of waterlogging tolerance of Brassica napus L.and provide reasonable improvement programs.There were six germination traits investigated on combining ability and heritability using complete diallel crossing method designed for 30 combinations from those six cultivars and lines.The traits included relative root length,stem length,fresh weight per plant,survival rate,electrical conductivity,and vigor index.After flooding treatment,the six traits of parents and F1 were analyzed.The general combining ability（GCA） and special combining ability（SCA） of germination traits were analyzed using Griffing I method.Among 30 cross combinations,the GCA was significantly different among six waterlogging resistance traits.The SCA of these traits was significantly different except the SCA of electrical conductivity.As a representative trait of waterlogging tolerance in rapeseed,relative vigor index had the highest narrow heritability and relatively low broad heritability.The cultivars Zhongshuang 9 and P79 had higher tolerance potential to waterlogging stress.It can be concluded that combining ability and genetic effects of relative vigor index during germination stage could be used to identify the waterlogging tolerance of rapeseed in breeding program.

Waterlogging stress in cotton:Damage, adaptability, alleviation strategies, and mechanisms

Over the last few decades,waterlogging stress has increasingly threatened global cotton production.Waterlogging results in reduced soil oxygen,impairing the growth and development of this valuable crop and often resulting in severe yield loss or crop failure.However,as cotton has an indeterminate growth habit,it is able to adapt to waterlogging stress by activating three mechanisms:the escape,quiescence,and self-regulating compensation mechanisms.The escape mechanism includes accelerated growth,formation of adventitious roots,and production of aerenchyma.The quiescence mechanism involves reduced biomass accumulation and energy dissipation via physiological,biochemical,and molecular events.The self-regulation compensation mechanism allows plants to exploit their indeterminate growth habit and compensatory growth ability by accelerating growth and development following relief from waterlogging stress.We review how the growth and development of cotton is impaired by waterlogging,focusing on the three strategies associated with tolerance and adaptation to the stress.We discuss agronomic measures and prospects for mitigating the adverse effects of waterlogging stress.

Effects of Waterlogging on Photosynthesis and Antioxidant Enzyme Activities of Six Barley Genotypes with Different Waterlogging Tolerance

A field experiment was carried out to study genotypic difference in the effect of waterlogging on photosynthesis, chlorophyll content and antioxidative enzyme activities in barley. Waterlogging caused a rapid decline in net photosynthetic rate (Pn) and stomatal conductance (gs), and little change in chlorophyll content during early days of the treatment. A dramatic increase in malondialdehyde (MDA) content, superoxide dismutase (SOD) and peroxidase (POD) in waterlogged plants in the early days of the experiment was found, indicating the occurrence of oxidative stress in barley plants exposed to waterlogging. There was a highly significant difference in the changed extent of all these parameters among genotypes. Franklin and Yongjiahong Liuleng Damai, which were relatively sensitive to waterlogging in terms of growth, photosynthesis and chlorophyll content, accumulated much more MDA than the other two relatively tolerant genotypes (93-3143 and QS). After removal of waterlogging, the genotypic difference became much greater in recovering of these examined parameters. Yongjiahong Liuleng Damai showed higher recovery, while Franklin only recovered to 50% of the control at the 14 day after waterlogging removal. It may be concluded that it is the difference in anti-oxidative stress caused by waterlogging that account for the major difference in photosynthesis among barley genotypes.

Differences in Chlorophyll Fluorescence Parameters, Yield and Its Components Between Different Genotypes of Wheat Under Waterlogging Conditions at Anthesis

Waterlogging is one of the most factors limiting wheat production in the middle and lower reaches of the Yangtze River Plain of China,especially in the middle and late stages of wheat.Wheat varieties‘Jingmai102’(JM102)and‘Yangmai158’(YM158)were planted to study the dynamic changes of photosynthetic characteristics in flag leaf and the influence of waterlogging at anthesis on the yield and components and dry matter accumulation and remobilization of winter wheat in above ground.The results showed that the SPAD values slightly increased at 1 day after anthesis(d),and then kept decreasing with the increase of waterlogging time.The decrease in SPAD value was more remarkably in YM158 than that in JM102.As for the chlorophyll fluorescence parameters,the photochemical efficiency(Fv/Fm),potential activity(Fv/Fo)of photosystem II,and electronic transmission(Fm/Fo)on photosystem II increased first and then decreased with the increase of waterlogging days after anthesis.The quantum ratio of heat dissipation(Fo/Fm)had a tendency opposite to that of Fv/Fm,and the change range of JM102 was lower than that of YM158.For the grain yield and components,waterlogging at anthesis decreased the dry weight of single stem,grain yield,1 000-kernel weight,spikelet per panicle,and harvest index,and the reduction of JM102 was smaller than that of YM158.As for the accumulation and remobilization of dry matter,the accumulation of dry matter after anthesis decreased significantly under waterlogging condition(WL),and the reduction of JM102 was smaller than that of YM158.In conclusion,waterlogging at anthesis significantly affected the photosynthetic characteristics,yield and components in both varieties,but different varieties exhibited different tolerances to waterlogging stress and YM158 was more sensitive to water stress than JM102.

Unraveling waterlogging tolerance-related traits with QTL analysis in reciprocal intervarietal introgression lines using genotyping by sequencing in rapeseed(Brassica napus L.)

Soil waterlogging is a major environmental stress that suppresses the growth and productivity of rapeseed(Brassica napus L.).Natural genetic variations in waterlogging tolerance(WT)were observed but no QTL mapping has been done for WT related traits in rapeseed.In this study,QTL associated with three WT related traits including relative root length(RRL),relative hypocotyl length(RHL)and relative fresh weight(RFW)were dissected using a set of reciprocal introgression lines(ILs)derived from the cross GH01×ZS9,which showed significant difference in WT.Genotyping-by-sequencing(GBS)of the populations were performed,totally 1468 and 1450 binned SNPs were identified for GIL(GH01 as the recurrent parent)and ZIL(ZS9 as the recurrent parent)population,respectively.A total of 66 distinct QTLs for WT at the seedling establishment stage including 31 for RRL,17 for RHL and 18 for RFW were detected.Among the 66 QTLs,20(29.4%)QTLs were detected in both genetic backgrounds and then they were integrated into six QTL clusters,which can be targeted in rapeseed breeding for improvement of WT through marker-assisted selection(MAS).Based on the physical positions of SNPs and the functional annotation of the Arabidopsis thaliana genome,56 genes within the six QTL cluster regions were selected as preliminary candidate genes,then the resequencing and transcriptome information about parents were applied to narrow the extent of candidate genes.Twelve genes were determined as candidates for the six QTL clusters,some of them involved in RNA/protein degradation,most of them involved in oxidation-reduction process.These findings provided genetic resources,candidate genes to address the urgent demand of improving WT in rapeseed breeding.

Waterlogging risk assessment based on self-organizing map(SOM)artificial neural networks:a case study of an urban storm in Beijing

Due to rapid urbanization, waterlogging induced by torrential rainfall has become a global concern and a potential risk affecting urban habitant's safety. Widespread waterlogging disasters haveoccurred almost annuallyinthe urban area of Beijing, the capital of China. Based on a selforganizing map(SOM) artificial neural network(ANN), a graded waterlogging risk assessment was conducted on 56 low-lying points in Beijing, China. Social risk factors, such as Gross domestic product(GDP), population density, and traffic congestion, were utilized as input datasets in this study. The results indicate that SOM-ANNis suitable for automatically and quantitatively assessing risks associated with waterlogging. The greatest advantage of SOM-ANN in the assessment of waterlogging risk is that a priori knowledge about classification categories and assessment indicator weights is not needed. As a result, SOM-ANN can effectively overcome interference from subjective factors,producing classification results that are more objective and accurate. In this paper, the risk level of waterlogging in Beijing was divided into five grades. The points that were assigned risk grades of IV or Vwere located mainly in the districts of Chaoyang, Haidian, Xicheng, and Dongcheng.

THE STUDIES ON WATERLOGGING DAMAGE IN JIANGHAN PLAIN USING DEM

The waterlogging damage is the pit to agriculture in plain and lake region. This damage is related to the groundwater, ancient lakes, soil, land use and negative landforms, the conventional technique is adopted to the study of this damage. In this paper, we suggest a new technical method, the technique is based on DEM, to study this problem. The DEM is developed on the ECLIPSE MV/ 10000 AOS/ VS system, and the estimation model of waterlogging loss is built on the historical data of the test region in Jianghan Plain. and then, the rice waterlogging loss of test region is estimated by them.