Posted on Authorea 27 Apr 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158800944.47703853 | This a preprint and has not been peer reviewed. Data may be preliminary. ocliaeteete pce nsln agnllns(un ta. 05.Pat rwn nsln soils saline CO in growing net Plants in 2015). decrease al., The et (Quinn crucial lands damage. is marginal mechanisms salt-induced saline Na adaptive on excess molecular the species and prevent tree 2020a). hormonal these Attipalli, saline physiological, cultivate & towards of to gain Marriboina economic 2016; understanding also al., of comprehensive but et are A lands Hanin species marginal tree 2013; of saline biofuel al., complexity to fixing et sustenance nitrogen genetic (Samuel introduced for important were lands the species only economically plant not by certain several spatio-temporal and importance issue, and hampered this lands productivity immense plant address saline greatly the To crop the of the climate 2019). is diversity rehabilitate al., extend Global genetic to attempts et to and (Morton these salinity made physiological 2019). soil on being of of (Hossain, heterogeneity depends are progress largely high attempts 2050 which However, land date, of year tolerance, cultivable To salt by because of lands. availability 2019). year lost the al., saline every be limiting et on by further (Raza will are lands production lands irrigation crop marginal arable excessive for saline and of population becoming closely 50% increasing is Approximately are nearly change, which 2020a). land and Attipalli, production cultivable & levels and Marriboina of salinity growth 2018; hectares plant al., et limit million (Shahid to 1.5 degradation constraint land environmental arable with major associated a is salinization Soil these salinity Collectively, conferring Pongamia. INTRODUCTION adaptations in metabolic homeostasis and ROS molecular and structural, ionic physiological, integrated the expression Pongamia. an differential regulate to that on genes showed tolerance insights antioxidant data our new as maintaining level, shed well in molecular aid results the as to At genes acids) Pongamia. organic transporter and to and JAs tolerance of polyols Na ABA, salt (sugars, including apoplasmic phytohormones conferring metabolites hormones and as that compatible that balance such selective demonstrated revealed osmotic with studies strategies interaction Correlation also plant adaptive positive Pongamia. in salt study a of adjustments the showed Our roots induced SA promoting and salinity leaves in observed. level, in role lignification were physiological crucial wall At patterns cell play days. leaves accumulation ABA 8 in and ion investigation for variations JAs present and NaCl) molecular including The (3% exchange and level elusive. physiological gas salinity are with leaf sea environment correlation morphology, at saline in Pongamia to responses of mechanisms metabolic roots adaptive and and the hormonal crop, in tree alterations semiarid Although tolerant describes lands. salt cultivable of a loss be and productivity, to plant in losses significant results stress Salinity Abstract 2020 27, April 3 2 1 Sharma Rameshwar Marriboina Sureshbabu in high tolerance of salinity insights provides network hormone-metabolite Systematic ffiito o available not Research Affiliation Advanced of Institute Indian Hyderabad of University + 3 o ipsto ntelae nodrt rtc h htsnhtcmcieyfrom machinery photosynthetic the protect to order in leaves the in disposition ion n tial Reddy Attipalli and , 1 ai Sharma Kapil , ogmapinnata Pongamia 1 eahe Sengupta Debashree , 2 1 siiainrt n piu unu il fPSII of yield quantum optimum and rate assimilation 1 L)pierre (L.) 2 nrp Yadavalli Anurupa , ogmapinnata Pongamia + eusrto and sequestration 1 sreported is , Posted on Authorea 27 Apr 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158800944.47703853 | This a preprint and has not been peer reviewed. Data may be preliminary. oe omn-eaoieascae inligptwy oudrtn ihslnt oeac mechanisms as well tolerance as salinity interactions high mechanisms hormone-metabolite understand the tolerant to for stress pathways evidence in an understand signalling provides to associated study dis- present hormone-metabolite crucial the The novel very for 2017). are opportunities al., The which new et create conditions. 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(2015), vacuolar and expression membrane as causes regulates V-PPase, such compartments ABA NHX1, transporters celluar are depolarization membrane and stress membrane which plasma cell salinity salt-induced and the genes alleviates the in SA coding salts to with protein of Counteract pre-treated several deposition Plant towards Excessive the 2020). plant 2018). regulate al., of survivability et positively Na (Wang the decreasing SA tolerance enhance by salt to and the plant cell regulate JA for 2019). the positively Additionally, Husen, responsible of can & status (Siddiqi JAs al., antioxidant stress. stress Further, regulates et salt osmotic during anthocyanin together (Gupta 2020). movement ABA directed cell Baek, understood guard and jasmonates & clearly the JA adapted regulate (Ali not together plants effects still level, damage, negative ABA are endogenous ROS its interactions induced mitigate their However, salinity to but species against accumulation rice. plants, plant combat in on on To tolerance depending imposed mostly salt 2017). stress are the significant of phytohormones improving a type between cause in interactions and can antagonistic role levels and significant synergistic SA a the endogenous played in phytohormones H raise (Liu among and the capacity harmony Na superoxide stress, photosynthetic of and salt marinating levels Upon ROS and elevated senescence the 2017). leaf al., in delaying et improves reduction by cytokinin Gloan of 2012; species accumulation such al., crop endogenous process cytokinins et in induced developmental and Salt tolerance and Auxins salt differentiation. growth 2020). and the various al., elongation excess regulate et division, prevent to (Zelm cell to interacts stress as phytohormones closure salinity promoting under stomatal extreme growth growth causes under regulating root are growth ABA and regulate its growth as and 2007). sustaining plants such evaporation (Tuteja, for in water phytohormones critical role responses phytohormone, several crucial signalling induced plays produces downstream stress which plant well-known numerous SA, is stress, Ca and ABA several salt IBA activating development. conditions. to IAA, through saline and response zeatin, toxicity growth In MeJA, sodium plant JA, of 2019). in ABA, amelioration (Thor, role in channels important (Ca role and an ion essential plays genes Calcium an and stress. plays messenger drought also second induced It intracellular salt an under as performance known leaf the substantiate might (Fv/Fm) ogmapinnata Pongamia + rnpr n yicesn H increasing by and transport . + APs up Fkd n ak,20) codn oSazde al., et Shahzad to According 2006). Tanka, and (Fukuda pumps -ATPase + cuuainars h ln,weesS ecetpatpoue an produced plant deficient SA whereas plant, the across accumulation 2 O 2 Yn ta. 04.Acrigt ao ta. 21) h perfect the (2014), al., et Sahoo to According 2004). al., et (Yang + APs ciiy(aaanne l,21;Gabe al., et Gharb 2013; al., et (Jayakannan activity -ATPase 2 + + iogncprpopaae aulrH vacuolar pyrophosphatase, -inorganic APs up,CX n Cswr involved were CCXs and CHXs pumps, -ATPase + ue nadoto h el(rgig tal., et (Dragwidge cell the of out and in fluxes + o paea the at uptake ion 2+ + responsive -ATPase, 2+ is ) Posted on Authorea 27 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158800944.47703853 — This a preprint and has not been peer reviewed. Data may be preliminary. ouini . OSbffroengta 4 glutaraldehyde at 2.5% overnight with buffer incubated and MOPS sections M cm 0.1 1 into in segmented solution were roots and leaves collected Freshly Chloride method Australia). 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CO (Qubit ppm CO 350 min after with taken ml. flushed 50 were continuously of chamber rate leaf flow 1600 to a attached of All at (PAR) (A113), Systems). radiation source active light (Qubit photosynthetic LED CO650 by saturating Q-box a 2 conditions: flow-through following expended the the fully with on measured performed were was measurements capacity photosynthetic -80 parameters net at fluorescence Leaf stored chlorophyll and and nitrogen exchange liquid in gas 4 frozen Leaf 1, flash control 2020b). 70 of intervals, of at Attipalli, interval weights time drying Dry & an each harvest. exposed after at (Marriboina after at determined and harvested day immediately harvested were measured chosen were per tissues were were Plants plant NaCl roots and solution. treated 30) mM leaves and Hoagland’s to 100 of weights water fresh of 20 Fresh days. 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Data may be preliminary. rudt odrwt iudntoe.Apoiaey10m fpwee isewssseddi 500 in suspended was tissue powdered finely of were plants mg salt-treated 100 and Approximately control (2010) nitrogen. of al., et liquid roots Pan with and μ from leaves powder protocol harvested a to Freshly using ground LC-MS by modifications. quantified some were with phytohormones of levels endogenous The analysis LC-MS nlhaiga 9@ o i.Teflwrt fcrirgs(eimgs a dutdt . l min ml. 1.5 to adjusted was gas) (helium gas carrier s. of scans/ rate 2 flow at The 600 min 1 at min. 5@C of maintained 5 by volume followed for were A min, 290@C 0.25 temperatures 5 at and for injection heating 70@C diameter of final at internal source heating mm isothermal and 0.25 tions: interference with ion, column The Rxi-5ms 200 Cor- USA). m (LECO 30 system (Restek, GCXGC-TOF-MS with thickness LECO-PEGASUS equipped system a USA) on poration, analysed was sample derivatized The -80 in kept Duran) were Schott samples (GL-14 80 the Finally, in vial dissolved 30 min. glass was at 45 residue in for the transferred derivatization, drying For Vac was analysis. 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Data may be preliminary. orlto ae lseigaogmtbltsi ev n roots and GABA leave and in metabolism metabolites acid among metabolites points. clustering amino other all-time based synthesis, Several across Correlation 8DAS. acid plants at organic treated plants acid, of treated NaCl to leaves fatty NaCl glycerol, mM mM in related to Leu, 500 Val, 500 shunt relating metabolites and pinitol, and several marginally pTag, 300 300 pGlc, accumulated increased of Fru, leaves roots also of includeMan, Pongamia in which roots similar benzoate addition, acids, Importantly, in and organic 8DAS. In threonate fold and and 8DAS. polyols ˜12 4 and synthesis, 1, by wall 4 at cell up-regulation 1, plants significant at treated showed plants NaCl levels treated mM mannitol 500 methyl-succinate leaves, and and 300 metabolic to phytol in palimitate, various acid laurate, increase and fatty to myristate, accumulated significant and NAG, also polyol showed acid related mannitol, plants amino Gal, metabolism, treated polyol, metabolites includes alcohol, salt which acid carbohydrate primary of metabolism, carbohydrates, amino roots cell-wall of Additionally, to metabolism, related set 6). metabolites carbohydrate various Figure a (Supplementary metabolism, accumulated metabolism acid acid roots fatty amino several amino points. as stress, Additionally, all-time synthesis, such salt 8DAS. across acid pathways plants at to organic treated plants pathway, response of treated glycerol leaves In in NaCl to several shunt mM related accumulated GABA mM fumarate, in 500 Pongamia and Leu, marginally 500 and pFru, metabolism time, pGlc, increased 300 acid and Xyl, treatment were of Gal, 300 increasing leaves include metabolites 8DAS. of which With in other and cycle, leaves 8DAS. glutarate 4 TCA in and 1, and and synthesis at fold succinate wall 4 plants cell ˜12 1, to treated related by NaCl at metabolites mM up-regulation plants 500 various significant treated and accumulated coumarate showed 300 NaCl leaves butanoate, levels both Val, exposure, in Phe, mannitol stress myo-inositol, increase Importantly, and mannitol, significant salt polyol pinitol, showed Upon metabolic Man, myristate metabolism, Fru, various and acid Suc, to 5). organic includes related Figure which metabolism, metabolites (Supplementary metabolites, carbohydrate primary metabolism of metabolism, acid set acid a fatty polyol amino accumulated cyclitol/ as leaves 6 such stress, pathways metabolism, salinity metabolites. acid to miscellaneous amino response other 15 In gas 10 metabolism, related and using are acid metabolism by which metabolites organic acid 2), quantified 71 Table fatty 22 (Supplementary of were metabolism,6 metabolism, roots total conditions and carbohydrate A 1) stress 12 Table (Supplementary 6). to salt leaves and both under 5 in Pongamia identified Figures were (Supplementary in at (GC-MS) NaCl metabolites spectrometry mM of chromatography-mass 500 levels of and relative roots 300 and The and leaves of (JA in roots Jasmonates profiling in 2D-F). metabolite hormones with (Figure Temporal other correlation points significant all-time with 2G-L). showed across interaction (Figure JA plants significant time each points Similarly, treated over showed all-time of 2A-C). NaCl IAA SA responses (Figure mM and and interactive plants 500 JAs MeJA) the treated of between was show leaves NaCl correlation point to in mM significant time matrix MeJA observed 300 stress We cluster of Ac- Each correlation leaves stress. a in stress. salt time. of salt under treatment form hormone under stress the individual interactions on in mM hormonal based 500 individually explore performed to grouped to were response essential studies In 8DAS. are correlation 1G). at hormones cordingly, (Figure ˜3.0-fold by among respectively by increased plants mM increase studies were treated significant 300 Correlation 2.0-fold levels showed NaCl and IAA of levels mM ˜3.3 and IBA roots showed 300 zeatin only levels of in 8DAS, stress, JA roots At ˜3.7-fold NaCl and 1F). in by MeJA (Figure 2.3-fold treatment, increase respectively and NaCl 4DAS significant ˜2.2 mM plants at showed 500 treated roots to zeatin NaCl in response mM only up-regulation and In plants 500 4DAS, NaCl concentration treated in plants. At NaCl mM up-regulation treated salt mM 1E). significant NaCl 300 300 with showed (Figure of varied of levels significant roots 1DAS ABA leaves levels showed in and at significantly phytohormone in JA zeatin increased Similarly, the enhanced levels and MeJA 1DAS. significantly roots, MeJA and at were IBA in IAA, Zeatin, levels Moreover, zeatin, time. JA 8DAS. treatment as and at such ABA plants Hormones IAA, treated significantly. 4DAS. change at not up-regulation did levels SA 7 ogmapinnata Pongamia .pinnata P. ne atstress salt under ne atstress salt under Posted on Authorea 27 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158800944.47703853 — This a preprint and has not been peer reviewed. Data may be preliminary. at cd t1A Fgr D.Icesditrcin eeas bevdbtenhroe n metabo- and hormones between observed observe also were could interactions we Increased plants, 5D). treated (Figure MeJA:IBA:zeatin:JA:IAA::carbohydrates, 1DAS NaCl other at including mM and acids and 500 metabolites StA, fatty JA:IBA:IAA:SA:zeatin::polyols Asn), of acids, and ABA:IBA:IAA:zeatin::amino Asp, leaves acids, hormones (Hse, In JA:MeJA:IAA:zeatin::organic metabolites group between 5C). with acids interaction interaction (Figure amino significant CAA), Grl-g) more showed 5Hpip, were NAG, MA, JA (G3p, Pip, and (NA, metabolites metabo- ABA several group while with FA, acid correlation 300 and significant of organic showed leaves Man pTag, ABA in Gal, and 5B). acids Suc, JA (Figure amino 8DAS, 4DAS MytA, and At at carbohydrates lites 4DAS. at plants with at plants as treated association plants treated NaCl such treated strong showed NaCl mM NaCl metabolites also mM 300 mM with MeJA of and interaction 300 leaves JA significant in of addition, showed acids leaves In organic zeatin in and and MytA carbohydrates, MeJA acids, with IBA, JA, amino ABA, correlation Hormones including positive 5A). hormones of a leaves, (Figure showed In roots 1DAS JA metabolites. and and and zeatin hormones leaves between MeJA, interaction in the shows 5 metabolites Figure and hormones between treated pinnata clustering NaCl based mM 500 Correlation in points. 6). MytA:MT:Phytol) Table all-time Ba:Gal:Gri:PA, Supplementary at Oxopro:Asn:Thr:Phe:CAA:5Hpip:IBua:pFru:Thy, the HA:Pdo:Eta, plants in (pGlc:Fru) (e.g. Rib:AA, given other group each were LA:Asu:PI, pos- within carbohydrate abbreviations significantly strong Ala:ThA, and and associated a were names (SA:Oxa) plants, metabolites 4G-L; several group treated (Figure addition, acid NaCl significant In points mM organic time showed 500 the the levels In all within metabolite across 5). observed Table the was with Supplementary correlation of the compared itive in most when given plants, were pronounced abbreviations treated PA:MytA:Gal:Gri:Met- names and less NaCl 4A-F; Glu:Oxopro:Thr:Phe:Hse:Ara:5Hpip:oC:Oxa:Pdo, (Figure were mM HA:MI:SA) Suc:GA:Fru, (e.g. metabolites Rib:Pyr:Glta:Eta, 300 SA:MT:Pi:Phytol:GlcNAc,pGlc:ThA:PI:pTag, other the of each among roots with correlations correlation In the datasets. dataset, leaf root the In each time within all interaction significant across a Asn:Hse:Pip) showed in Man:pFru:MI:MT:oC:IBu, points. points. metabolites pGlc:SA) Asp:Oxopro:Glu:5Hpip:CA:CAA:Grl-g, all-time significant several Xyl:SA:Pi, Additionally, at a (e.g. (Suc:Val, plants. Man:pFru) plants, other metabolites treated acid (Xyl:pGlc, treated other NaCl organic with NaCl group mM Leu:Phe), interaction mM carbohydrate 500 significant Asn:Hse, 500 and showed (Asp:Glu:Oxopro, In IBua:oC) also group Carbohydrates 5HPip:CA:CAA, plants. acids mannitol treated amino (e.g. myristate, the NaCl acids fructose, group within mM organic as a observed 300 with Additionally, such was of interaction metabolites plants. correlation leaves several strong treated in between showed NaCl myo-inositol observed levels mM and was 300 these correlation of group, correlation positive leaves significant acid strong the in a L; amino within GABA:MA) Although and correlations the Asn:Pip:5HPip, (MI:MT). 3I high polyols (Figure within (Asp:CA:NA:CAA, observed group and we observed plants, one 5HPip:Pip) not into treated CA:CAA:NA, was NaCl clustered (e.g. 8DAS mM group at 300 and acid 3H of NaCl (Figure were leaves organic mM group abbreviations In 500 one and into of 4). names clustered Table leaves 4DAS metabolite Supplementary at and the of NaCl mM 4) simplify order mM Table 500 500 J; to Supplementary of of and leaves order K; leaves 3G 4), Similarly, In Table (Figure Supplementary tables. group the 3). the in one Table given in into at Supplementary clustered separately treatment F; showed 1DAS treatment and NaCl were at NaCl (r[?]0.6) NaCl 3C mM mM correlations 300 (Figure 300 of strong metabolite group leaves of only and of another leaves 3) graphics, order Table into D; 3), Supplementary clustered E; and Table and 8DAS 3A Supplementary 3B (Figure time at (Figure the group treatment group a in a on into into based given al., clustered clustered groups 4DAS were 1DAS et was six at Sanchez abbreviations analysis into treatment 2011; Correlation and clustered NaCl al., were mM names (15,123) 4). values et 300 and NaCl correlation of (Moing 3 The leaves mM pair (Figures HAC e.g. 2011). 5041) 500 metabolite al., metabolite; stress, each (15,123), et 71 for salt NaCl Lombardo X coefficients 2012; (71 mM to (30,246) correlation 8DAS 300 Pearson leaf and subjected tissues with treatments 4 different Pongamia performed 1, different two points two in of time include elements three metabolites 60,492 which and the total (30,246) a among root on relationship performed and was th ne atstress salt under 8 P. Posted on Authorea 27 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158800944.47703853 — This a preprint and has not been peer reviewed. Data may be preliminary. eepiesfrNX,NX,NX-ie O1 O1lk,H SOS1-like, and SOS1, (CDPK3) plants. NHX6-like, kinase3 treated NHX6, salt protein NHX3, roots Calcium-dependent for and decreased/ leaves was primers (CNGC5) plants. gene both Gene 5 treated CNGC17 in in channel of unchanged of induced level increased/ significantly ion roots expression were were nucleotide-gated the and levels levels Furthermore, leaves Cyclic CDPK32 8DAS. these and while 4DAS. both 4 plants in at 1, treated unchanged at only of plants leaves plants treated in of treated increased roots NaCl or decreased mM were PM-H 300 while levels plants, of treated roots salt PM-H of in leaves of in levels levels expression control The of plants. treated salt of nue nbt evsadroso attetdpat.W lomntrdepeso eeso he plasma three of levels expression monitored also We plants. treated salt (PM-H Vacuolar of ATPase roots plants. proton treated and membrane leaves NaCl 300 mM both of 500 in roots in induced in unchanged 1.9-fold proton remained and NaCl (V-H cation levels ˜2.5 subunit mM ATPaseB these ˜3.0, vacuolar 300 proton Under by though The ˜2.0-foldin increased plants, plants. significantly by treated treated levels significantly plants. NaCl these salt induced mM while treated of were 8DAS, laves levels roots salt at in (CCX1) and only of plants observed 1 leaves treated exchanger levels roots both was calcium expression in and increase cation in induced leaves ˜6.0-fold stress, changes marginally salt much and both were observe levels ˜6.0 in not (V-CHX1) ˜5.9, (CLC1) did 1 ˜3.7-fold as we exchanger 1 and However, well channel ˜2.0, respectively. as ˜3.9, chloride plants levels plants by treated of up-regulation 2 treated NaCl significant Sensitive) mM NaCl a Overly 500 showed mM unchanged (Salt levels were of 300 levels SOS SOS3 these of The plants. while leaves 1DAS. plants, treated treated at in salt NaCl plants of mM treated roots 500 and the NaCl 300 (Figure ˜2.0-folds in mM Further, of 1DAS and leaves 300 6A). at in ˜3.6 of decrease plants by (Figure increase/ roots treated increased showed respectively significantly NaCl and 8DAS levels leaves mM at (HKT1:1) 500 both 1:1 300 NaCl 500 in in of transporter only mM of leaves affinity increased 500 leaves in high significantly Interestingly, unchanged and in was 6B). remained points expression ˜3.6-fold levels all-time gene these and NHX1 at while ˜2.6 the respectively, plants 4DAS by treated and up-regulated NaCl 1 significantly mM at plants was treated (NHX1) NaCl 1 mM exchanger 6). proton (Figure Sodium CDPK32) and (CDPK3 kinases protein dependent oass h ffc fslnt teso h eeepeso rfieo ogma emntrdteex- the monitored we V-H and Pongamia, SOS2 CCX1, of (SOS1, V-CHX1, profile components pathway expression CLC1, SOS gene HKT1:1, PM-H including the subunit, (NHX1, genes on salt-responsive transporters key stress SOS3), several salinity of of profile effect pression the leaves assess in To potential membrane In and transport of 5J-L). ion roots (Figure respectively. with and 8DAS associated plants genes at treated of acid expression ABA:MeJA::organic NaCl Differential namely carbohydrates, mM metabolites ABA:SA::cell-wall and 500 and hormones 4DAS, of between at was ABA:SA:JA:MeJA::organic roots acids interaction carbohydrates, significant JA:SA:IBA::cell-wall in 1 1DAS, plants, at day decreased treated acids from NaCl ABA:SA:IBA:zeatin::organic interactions mM progressed positive 500 salinity of the the roots As 8, 5I). day (JA:MeJA:ABA:zeatin::organic (Figure metabolites to JA:zeatin::polyols) and hormones acidand between noticed JA:IBA:zeatin::amino was at interaction acids, strong plants namely a 8DAS, metabolites (Fig- treated At and acids 5H). hormones NaCl ure MeJA:zeatin::fatty between and seen mM were MeJA:JA:zeatin::polyols interactions 300 carbohydrates, positive JA:ABA:ABA:SA:IBA::cell-wall more in 4DAS, At acids 5G). 8DAS (Figure MeJA:IBA::fatty 1DAS JA:SA:ABA:MeJA:IBA:zeatin::carbohydrates, at and including plants were acids metabolites treated and JA:MeJA:SA:ABA:IAA:SA::organic hormones NaCl between associations mM significant roots, 500 In of leaves in acids 5F). JA:ABA:MeJA:IBA:zeatin::fatty (Figure and JA:ABA::polyols, cell-wall acids acids, and fatty JA:ABA:MeJA:IBA:IAA::amino MeJA:JA:ABA:SA::carbohydrates pos- acids, namely significant JA:ABA:MeJA:SA::organic metabolites a and recorded carbohydrates, We hormones 5E). between (Figure correlation 4DAS itive at JA:MeJA:zeatin::polyols acids, acids, MeJA:IBA:IAA::amino JA:MeJA:IBA:zeatin::organic cell-wallcarbohydrates, and JA:MeJA:zeatin::carbohydrates namely lites .pinnata P. + Tae,PM-H ATPase1, + Tae uui)adV-H and subunit) ATPaseB + + Tae.,PM-H ATPase4.1, Tae sfrsPM-H isoforms ATPase) + + 9 Tae.-ie NC n NC7,adcalcium- and CNGC17), and CNGC5 ATPase4.1-like, Tae,41 n .-iegnswr iia othose to similar were genes 4.1-like and 4.1, ATPase1, + + + Tae.-ielvl eemrial increased marginally were levels ATPase4.1-like Tae uui xrsinlvl eeslightly were levels expression subunit ATPaseE Tae,41 .-iei ohlae n roots and leaves both in 4.1-like 4.1, ATPase1, + APs4 H -ATPase4, + Tae uui,V-H subunit, ATPaseB + APs4lk,CNGC17- -ATPase4-like, + ATPaseE Posted on Authorea 27 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158800944.47703853 — This a preprint and has not been peer reviewed. Data may be preliminary. h osqec fsl-nue hthroa itrac nlae n ot fPongamia. of roots and leaves 2018). (Wu, in toxicity disturbance sodium phytohormonal from salt-induced enzymes of cytosolic consequence of Na The protection for the facilitate matrix may cation-binding which as 2020a), Na Attipalli, serve that vaculoar primarily indicating and plants may wall treated apoplastic Na NaCl cell apoplastic mM the 500 in to and pectin Na Na 300 of of both apoplastic Na residues sequester wall in an effective cell time was an apoplastic/ treatment the there as study, with present act increased the roots In was intensity Pongamia 2020a). Attipalli, that & demonstrated (Marriboina ions studies fluorescence previous Our 2014). al., et Ca (Choi Intriguingly, conditions 2018). photosynthesizing Ca al., vaculoar and et Wu of Ca dividing 2018; trigger actively al., possible et Na on might Rahneshan increasing that effects 2016; Na with al., negative increased excessive et also the Peng for were 2016; alleviate sink al., and et potential (Baetz shoot as cells Na roots to act regarding the may translocation results that roots The their Na indicate the 2020a). of RRWC Attipalli, that levels unchanged High & suggesting and (Marriboina interesting. LRWC photosynthetic halophytes rather solution the reduced true are salt in accumulation The like process alterations NaCl) to insignificant 2017). able (˜3% and al., were NaCl morphology et mM leaf (Marriboina (500 sustained Pongamiamachinery salinity exhibiting in high by mechanisms mangroves to adaptive and tolerance salt significant possible exhibited the Pongamia are roots the in Ca tration parameters, photosynthesis Sustained and plants. 2 treated CAD1, expression all-time salt and the at However, of 2, DISCUSSION 8DAS. PAL1, roots increased and SOD, in 4 also Cu/ZnSOD1-like changed at FeSOD Cu/ZnSOD1, treatment slightly MnSOD, NaCl were treatment. including mM 6 CAT4 genes 500 salt genes, in other mM antioxidant and all plants the 500 of treated Among levels 8DAS. of salt and 11). mM plants 1DAS and 300 4, treated at in 10 NaCl 1, points 9B, only mM at 500 Figures plants up-regulated (Supplementary and treated significantly 300 respectively NaCl in was 8DAS mM up-regulated and showed substantially 500 4 were 6 1, in CAT4 at CAD levels as while expression down-regulated well the all-points, induction. significantly as were roots, significant at 4DAS levels In any and plants these 1 show while treated 8DAS, NaCl not NaCl at mM did mM treatment 300 NaCl CAD2) 500 only in mM and genes, and 300 to CAD1 strengthening 300 in PAL2, increase cell-wall 1 of significant (PAL1, the leaves from genes Among in treatment different 8DAS. other up-regulation till leaves, salt at the only strong In treatment up-regulation mM a significant stress stress. showed 300 showed prolonged isoforms salt POD under at SOD mM stress the declined up-regulation 500 treatment, but salt significant NaCl and 4DAS, the mM and 300 500 throughout gradual both under However, leaves showed in 8DAS. in DAS SODs 8 unchanged of at remained isoforms down-regulation expression showed CAT4 and genes treatment antioxidant 11). and the 10 Among PAL2, 9A, PAL1, Figures (POD, enzymes (Supplementary thickening CAD6) wall (CAT4, and cell enzymes antioxidant and CAD2, including Cu/ZnSOD1-like) genes and salt-responsive Cu/ZnSOD1, modifications key MnSOD, several wall FeSOD, of levels cell expression the and monitored We homeostasis ROS roots with primers and 8, associated leaves and genes in 7 Figures of Supplementary expression in 7). (given Differential Table plants Supplementary salt-treated the and control in given both were of roots and leaves in V-H and like, 2+ a on ob novdi rpgto fln itnesgaln ro-osot ne atstress salt under (root-to-shoot) signalling distance long of propagation in involved be to found was + ps xiie ihmlil adn atrsterepeso atr infiatyvaried significantly pattern expression their patterns banding multiple with exhibited ppase + eusrto Gnae ta. 02 arbiae l,21) h atrso high of patterns The 2017). al., et Marriboina 2012; al., et (Gonzalez sequestration .pinnata P. + + otnsmgtla olwrNa lower to lead might contents eusrto Gnae ta. 02 nwre l,21) h carboxylic The 2017). al., et Anower 2012; al., et (Gonzalez sequestration + eesi ohlae n ot fsl rae lns hc suggest which plants, treated salt of roots and leaves both in levels 2+ eevst mloaeteNa the ameliorate to reserves 2+ inligptwy n ppamcNa apoplasmic and pathways signalling 10 + n Cl and - + oswr eetdi ot ftetdplants treated of roots in detected were ions o otn ntectsl(arbia& (Marriboina cytosol the in content ion + n Cl and + oiiy(ae leh 2013). Plieth, & (Saleh toxicity - osdpsto,inhibiting deposition, ions + pcfi fluorescence specific + o,contributing ion, + n Cl and + 2+ seques- - levels ions + Posted on Authorea 27 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158800944.47703853 — This a preprint and has not been peer reviewed. Data may be preliminary. aiecniin,abi h inligmcaimi tl nla Dvnre l,21)(iue7). (Figure 2013) al., under et growth (Devinar plant improves unclear in which still favour plants, that is may treated revealed mechanism content salt studies signalling higher of ABA correlation the roots of reduced The stress in albeit from maintenance Further, SA salt 2015). conditions, transfer with prolonged al., of saline 2015). interaction et The ABA mitigation strong al., (Shi a in to et showed proteins 2015). helps ABA due (Shi aquaporin al., increase regulating genes be et transient by responsive RWC might (Shi while stomatal maintaining stress growth, et roots roots the the plant (Skorupa the in enhancing the limit apparatus from accumulation by impacts might exudation stomatal ABA negatively ABA which the levels in or plants ABA closing reduction shoot by treated observed to CER salt 2018) The root and al., of et rate 2019). leaves Yang transpiration al., in 2016; content, increased al., water et significantly guard conductance, Wang in were 2011; pathways levels al., signalling ABA et stress-induced (Munemasa the leaves triggering the by between 7). from interactions & closure (Figure The loss Tavallali stomatal plants. water stress 2015; treated induce preventing salt of al., leaves of may cells, et in effects ABA ABA Shahzad toxic with and the correlation 2013; Li JAs strong alleviate wall a al., 2016; level showed cell et JAs al., JAs and that Uddin increased et revealed uptake the Na 2005; Cao water that the enhanced al., 2014; lowering suggest growth, et by Baldwin, studies shoot (Kang Previous & reduced species Mitra 2019). JAs Karimi, crop 2014; of JA certain application al., transduction that in exogenous roots, et signal indicates An synthesis (Diallo in common results share 2017). stress However, The might al., 8DAS. salt and were et at interchangeable during plants. levels levels presumably Pongamia MeJA treated control are Conversely, in JAs to 8DAS. salt pathway of returned at of forms levels levels different leaves control these two to 2018). while both the restored al., 4DAS, in and et till 4DAS, Ishimaru high detected till maintained 2018; was low al., growth JAs maintained et salt-induced were Fattorini in (JA promoting levels 2014; JAs in increase al., defective involve with et significant showed might association (Cai which genes A strong stress plants, IBA a treated salt mM and showed salt during 300 IAA also integrity of of Auxins both tissue roots leaves in and and 2014). leaves in deficient al., in IAA plants et MeJA) the 2010). with (Spiess and play that interaction al., development also evident and et good may was (Tognetti significant growth showed levels It Pongamia plant showed IBA IBA in also plants. that enhanced changes levels treated revealed that or architectural NaCl studies IBA suggesting protective damage plants, correlation Likewise, stress-induced tissue integrity. treated addition, the salt 2012). tissue to In acquiring of levels al., and due in roots steady et cells role may and and (Kim a of leaves IBA trait stress status “stay-green” both and maintaining salt viable in IAA in increase under the help of overexpress- pigments suggest might levels chlorophyll Pongamia plants, clearly in increased of studies poplar levels The fluorescence IAA Transgenic that increased our indicating Therefore, 2012). However, points, tolerance. al., lysis. all-time salt et cell at Pongamia (Kim plants in senescence treated role leaf salt important of an leaves ing al., play in et increased might Jang 2016; significantly IAA al., were et Nitschke salt levels 1982; the IAA Kato, enhance RRWC (Ueda& maintained also 2D) well Figure may the by (Supplementary JAs plants. supported plants 2017). also and treated treated is NaCl of zeatin which roots uptake, mM between water under strong in 300 interaction survival enhance A plant to of synergistic promote 2016). roots roots The to in al., induced-vasculature and impact et negative leaves conditions. Melo JAs salt in 2014; saline substantiate under may JAs al., extreme growth cytokinin et and of Wu stress-induced zeatin maintain influence 2011; and interactive between RRWC of to al., The levels improved observed plant et increased plants also Nishiyama the Moreover, treated 2011; was for 2015). al., salt correlation beneficial al., mM et of et be (Ghanem 300 Fahad roots may conditions of 2014; ad correlation stress al., exposure leaves and et initial both hormones all- (Sahoo to in conditions all at due zeatin stress patterns in hormones salt correlation rise under all their The growth among and the leaves. observed profile in was hormone stress correlation in significant NaCl roots diversity and A significant leaves a both hormones, points. study, showed between time this plants interactions In treated the processes. salt developmental upon of and depends responses largely stress-adaptation regulate condit AtYUCCA6 eeascae ihicesdlvl faxn hwddlydclrpyldgaainand degradation chlorophyll delayed showed auxin, of levels increased with associated gene + n Cl and - osacmlto costepat(hha ta. 05.Creainstudies Correlation 2015). al., et (Shahzad plant the across accumulation ions 11 Posted on Authorea 27 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158800944.47703853 — This a preprint and has not been peer reviewed. Data may be preliminary. nlae,asrn orlto a bevdaogcroyrts at cd n oyl ntetdplants treated in polyols Na against and defense’ acids ‘osmolyte in line fatty first carbohydrates, studies as serve among may salinity observed metabolites these was for that correlation suggesting strong insights a new leaves, In provides to metabolites osmoticum source between observed intra-cellular 2018). carbon Pongmaia Kleczkowski, analysis we as providing & function Further, correlation Igamberdiev in to 2013; The essential known al., 2019). is et are al., (Bahieldin glycerol which et stress Moreover, pathway, salt Seifikalhor carbon TCA glycerol pathways. under respiratory 2019; maintain triglyceride of support induction al., to and and The metabolites cycle pyruvate glycolysis et shunt. TCA in free of GABA in (Che-Othman increase of carbon production the stress accumulation for of marginal the source salt the induction alternative in in marginal under an involve as a increase metabolism also serves these observed An may might we of shunt results, GABA levels Leu 2019). of these increase Increased al., and which with significant et Cys, production, Consistent plants. acid (Chen while Ala, treated salicylic cycle. stress as Thr, roots, salt salinity well acids of as and under biosynthesis amino roots leaves role lignin for in defensive both plants only play the cell in may to levels the beneficial benzoate ferulate be protect and and could to metabolites caffieate coumarate fluidity in in membrane observed increase was the stress an protect salt showed might Na under also from acids glycolysis organelle fatty maintain cellular saturated to and pyruvate of 2017). of accumulation al., the et production Increased Hossain limiting and involve 2005; by may pH, al., apparatus accumulation et cytoplasmic acid photosynthestic (Felle Lactic of condition the 2017). components regulation protecting active al., the in are (Hossainet glycolate in metabolites role and photo-inhibitory crucial serine toxic of a of accumulation ((Cao deposition play amino the macro-structures wall Interestingly, which cellular free cell of 2010). condition these photorespiration, of stability al., stress of as of biosynthesis et well salt levels Nasir including as under 2017; processes increased integrity al., growth protein biological The et membrane plant several 8DAS. the regulating support protect at by as to to plants tolerance components, uptake such treated to water acids contribute of and amino also roots nutrient and free and continue of leaves provide levels may in enhanced acids Phe source that and immediate and indicate Cys osmolytes also Thr, as 2016). serve profile wall may al., leaves metabolic call level et carbohydrates The Geilfus both secondary (Abdallaha in 2009; to in cell rise al., a contribute The recorded et for (Gilbert to 2019). were Pongamia energy al., known carbohydrates in of et 2003; are composition of Zhao wall also carbohydrates 2017; al., levels cell al., These but the High et et 8DAS. alters potential accumulation at (Abebe 2019). their osmotic plants while conditions al., synthesis, treated cellular Pongamia. et stress of in the Dumschott abiotic roots osmoticum regulates 2017; and major under only al., a protection as not et and scavenging serve Hossain alcohols leaves may ROS both carbohydrate mannitol in non-enzymatic that observed of provides suggest fold), levels 8DAS, (˜12 level at increased mannitol plants in The treated rise organic salt the acids, Interestingly, of amino roots carbohydrates, acids. including fatty level maintain and metabolite polyol, acids the acids, in fatty changes significant and exhibited stress to Pongamia root salt organic 2019). extensibility al., from cell under alcohols, et leaf correlation balance SA Shaki the carbohydrate positive osmotic of 2018; maintaining a cellular al., carbohydrates, transportation in revealed et help the cell-wall (Formentin studies might stress to Correlation plants Enhanced salt due treated under 2017). salt be growth al., of plant may al., leaves et better roots in et (Xu promote IAA in (Husen conditions and myo-inositol) levels stress SA regulate and SA salt between in also pinitol, in under SA and improvement (mannitol, in pathways reduction shoot levels an rise signalling polyols to observed The these necrosis showed carbohydrate The inhibiting 2018). while plants accumulating by al., 4DAS, by 2018). on et injury at turgor Husen salt SA 2013; from high leaf of al., leaves the maintained et the application (Jayakannan and protect stress Exogenous 1DAS might salt levels 8DAS. under at homeostasis at low ROS levels little and LRWC maintained control were to roots returned in levels SA + oiiyadinlaae(h ta. 05 tkje l,21) u results Our 2019). al., et Atikij 2015; al., et (Shu leakage ion and toxicity 12 + o soi imbalance, osmotic ion β- l,Vl e,Ala, Leu, Val, Ala, Posted on Authorea 27 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158800944.47703853 — This a preprint and has not been peer reviewed. Data may be preliminary. atrso O1adSS-iegnscreae ihNa with correlated genes SOS1-like and inclu- genes SOS1 transporter of salt-responsive patterns of PM-H expression NHXs, specific SOS1, tissue exhibited ding Pongamia under genes study, balance present transporter the osmotic ion In and of expression energy of in carbon association B alteration under increased of 2017; Salinity-induced regulation The source al., the pH et immediate 2020a). cell- with (Assaha as Attipalli, provide and conditions well decreased & may hormones stress as were Marriboina between acids salinity permeability 2017; interactions correlation organic water al., high number with the et The the hormones enhance (Marriboina plants. contrast, may conditions treated In tress acids time NaCl salt organic treatment mM plants. and increasing 500 carbohydrates treated of with wall NaCl 2014; roots increased in mM al., metabolites time et 300 and treatment of (Sahoo hormones and stress between roots hormones salinity interactions in between extreme of interaction under number increased survive The the al., to that et Pongamia suggest 2018). by (Sharma for al., results growth stress et beneficial salt-induced The salt Formentin promoting under be 2019). in balance may treatment involved nitrogen Husen, salinity metabolites be and the & may carbon with Siddiqi acids homeostasis, acids 2018; amino ion amino showed balance, and and soybean 2017). osmotic acids acids of Latif, regulating organic organic & shoots with (Mohamed that on stress interaction MeJA suggesting drought strong of time under showed application growth exogenous also maintain to The JA SFAs amino Further, wall 2018). N-terminal cell processes al., of biological the et accumulation several stresses Majeran to increased of environmental 2016; various regulation acid) and under al., in stability involved (fatty et S-acylation protein be (Boyle lipid as and could associations such modification of protein-membrane lipid modifications homeostasis, link The redox lipid irreversible proteins. protein of the the residue to acid favour correspond may for which acids insights fatty N-myristoylation, with new association provides Hormones metabolites and phytohormones studies between salinity analysis correlation The fNX a nhgduo 0 Msl tesi ohlae n ot fPnai,wieteewas there while Pongamia, of roots and leaves expression both of the in tolerance that stress salt observed salt in we study, role mM crucial this 300 their In upon suggest 2017). unchaged roots al., was and et leaves NHX1 (Marriboina both of pathway the SOS Na SOS3in significantly through and apoplastic increased Pongamia SOS2 to genes of contribute SOS1-like and expression may SOS1 tial which of roots, levels salt-treated expression the in Further, Na region. low apoplastic for xylem/ responsible are genes SOS1 of uptake increase the by provide may raised polyols imbalance of osmotic association the positive mitigate The the to 7). enhances level (lignin (Figure synthesis it cellular Na 2018) barrier at where al., wall osmoticum defense’, et cell continuous Yang barrier and a 2018; detoxification wall ROS al., cell these as et line with and such (Liu pathways metabolites ‘antioxidant as biosynthesis) regulating secondary fluidity of by serve of membrane line salinity correlation may to and second enhanced tolerance acids The potential as fatty osmotic serves 2015). al., pH, could and et cellular metabolites carbohydrates Guo regulating 2002; acids, defense’, Murata, & organic osmolyte (Sakamoto and between homeostasis correlation ‘ion positive 2010;Naliwajski of energy the al., carbon et roots, (Nasir immediate In Pongamia showed provide acids in ‘quick amino and stress and line salt-induced carbohydrates cycle correlation, third between ameliorate Sk2018). association TCA positive may lodowska, & as The which The the serve 2017). balance, for al., might C:N 7). et maintain role acids (Figure (Zhang stress 2015) organic anaplerotic salt al., under and the an source maintain et acids suggesting to carbohydrates, (Guo amino defense’ defense’ between stress homeostasis imbalance polyols, interaction energy salt ‘ion osmotic positive carbohydrates, line under the from second between balance Further, as membrane observed ion serve 7). cell and could (Figure polyols pH and 2015) and intracellular cell al., cycle the et TCA of Conde of intermediates 2013; integrity al., functional et and (Gao structural the protecting + osit h aul Saae l,2015). al., et (Slama vacuole the into ions + APss V-H -ATPases, + eesi h evso ogmab nraigNa increasing by Pongamia of leaves the in levels + APss NC,adohrtasotrgns h expression The genes. transporter other and CNGCs, -ATPases, h ta. 08 ag&Go 2018). 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Data may be preliminary. altetdrosmgtpri h ln opeev h elwl nert ne ihsl environment salt high under integrity wall mM cell 500 the in PAL2 preserve and to PAL1 plant both the levels of expression permit levels the might expression addition, PAL2, increased PAL1, roots In include, The treated 2012). which maintaining CAD6. monitored al., NaCl in were and et genes role CAD2, (Kim biosynthesis correlated CAD1-like, major stress lignin well and salt CAD1, a were modifying under play cell-wall roots wall salt peroxidases other cell and several high apoplastic of the leaves under classIII of in system integrity The antioxidant POD help structural depositions. the of the might lignin of expression specificity respective roots Increased tissue the in 2017). and with H levels al., dynamic of et expression the (AlHassan reflect levels CAT4 environment may higher higher leaves Also, the the (2010) in 7). reduce Tuteja, CAT4 (Figure to & biosynthesis Gill plant cellular lignin to the the of before According through levels much salinity. wall expression stress high oxidative higher cell under of observed function onset the induced H the its study, catalase at the ROS initiates this plants (SOD), the cascade the In dismutase protect signaling combat 2004). superoxide might antioxidant To al., isoforms as et SOD 2016). such different (Reddy al., the enzymes (POD) antioxidant et peroxidase several (AbdElgawad and induces (CAT) (ROS) plant species damage oxygen cellular reactive generation 2015). al., and et Saand genes Na 2013; antioxidant Excess al., of et (Wang expression stress Ca in salt induce alteration may of Salinity-induced CNGC17-like effects and negative CNGC17 the 2017). CNGC5, mitigate al., of to Ca et expression reponses Differential Liu intracellular 2018). 2014; of al., al., regulation et et Corso in Qi involved 2014; al., may Na et CCX1 vacuolar homeostasis, (Guan ion of conditions cellular improving experssion saline by under Enhanced growth plant osmoregulation in and involve balance may V-CHX1 pH of 2020a). levels Attipalli, expression increased & Na An membrane Marriboina of vacuolar 2018; of depostion al., depolarization excess et the PM-H by control (Graus may exposure of generated plants salt treated levels is salt V-H under which expression of of isforms leaves potential, expression the al., in these increased subunit The in of et stress. E diffrences salinity modifications and (Olfatmiri ATPaseB under post-translational The growth conditions promote and/or 2016). could stress toconfigurational which al., stress, salt due et be under Shabala could homeostasis 2016; isoforms ion al., and et potential 2014;Falhof not transmembrane may CLC1 balance, that pH indicates Na CLC1 of Na promote levels of PM-H Induction may expression retrevial The 2018). roots Cl 2016). the al., the al., and regulate in et et leaves involved HKT1:1genemay Ali Zhang be 2007; of both 2017; al., expression in et al., marginal (Davenport stress et xylem the salt An time, 2013; of treatment al., stress imposition et Na initial Hill tolimit upon plants. 2012; exclusion stressed expression al., Na salt HKT1:1 et of Na of (Munns of exclusion root-to-shoot roots phloem mediate and shoot-to-rootand can into leaves proteins translocating stream both transporter by in family roots trend HKT et and that similar (Bassil suggested followed stress studies HKT1:1 salt Previous of under study,expression development this and roots growth and In 2018). leaves plant al., both homeostasis, et in ion NHX6-like Dragwidge nd in 2018; NHX6 al., roles NHX3, as possible such Na their isoforms vacuolar suggests NHXs for other isofoms of expresssion NHXs differential other of involvement levels Na ting low sequester Na the to the with Na experssion mitigate correlates vacuolar NHX1 which to induce roots, might vacuole Pongmia and the NaCl), leaves mM both (500 in concentration stress NaCl Na mM of 500 in induction significant + 2 + O APs aiypmspaacuilrl nipoigtesl oeac ymitigteintracellular the maintaing by tolerance salt the improving in role crucial playa pumps family -ATPase ursneitniyadNa and intensity fluoresence 2 + rdcdtruhSDatvt,poie nadtoa datg otepatb strengthening by plant the to advantage additional an provides activity, SOD through produced + + cuuaini h ln assinciblnewihrslsi o oiiy xdtv stress, oxidative toxicity, ion in results which imbalance ionic causes plant the in accumulation eusrto,RSadinhmotssudrsl tes(oge l,21;L ta. 2016; al., et Li 2014; al., et (Yong stress salt under homeostasis ion and ROS sequestration, eusrto.I otat H1lvl eenffce tita moto fsrs,sugges- stress, of impostion intial at wereunaffected levels NHX1 contrast, In sequestration. + - aulrsqetto Wie l,2016). al., et (Wei sequestation vacuolar oct ntersetv ise(hn ta. 08.Hwvr ihicesn salt increasing with However, 2018). al., et tissues(Zhang respective the in toxcity + oiiy iial,i ot,idcdNX xrsinmyidct the indicate may expression NHX1 induced roots, in Similarly, toxicity. + otn aai evso 0 MNC rae lns thg salt high At plants. treated NaCl mM 300 of leaves in data content 2 O 2 ne ihsl odtos h oe xrsinlvl of levels expression lower The conditions. salt high under 14 + + osi evso ogmauo rlne salt prolonged upon Pongamia of leaves in ions eusrto tita tgso atsrs.The stress. salt of stages intial at sequestration + eieyb ihran Na withdrwaing by delivery 2+ ees hc a epin help may which levels, + + PaeadV-H and -PPase osrpdyinto rapidly ions + + rmxylem from rmleaves from + 2+ + rmthe from -ATPase derived + + - Posted on Authorea 27 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158800944.47703853 — This a preprint and has not been peer reviewed. Data may be preliminary. infiat (P * significant) mean the represent bars Na of Cl ratios the (E) Ca Na in (%), (F) content (RWC) (G) plants, water plants plants, relative salt-treated salt-treated (D) and and ( content, control control chlorophyll (CER) of of total rate roots (C) exchange and (FV/Fm), carbon leaves PSII of of Responses efficiency (A) photochemical of contents ion and S2. Figure S1. the Figure supervised A.R.R. paper; bioin- the performed wrote INFORMATION discussion. S.M. SUPPORTING D.S. the and and in K.S. S.M., data; A.D.Y., helped R.P.S. the research; and analysed the research D.S. performed and K.S. S.M. and analyses; S.M. formatics research; the designed S.M. of Department CONTRIBUTION from A.R.R. AUTHOR (BT/PR-12024/BCE/8/1097/2014) to grant India project of Govt. the (DBT), by Biotechnology supported was research This declare. to FUNDING interest of conflicts no have fellowship. Authors for India Delhi, INTEREST New OF (UGC), Commission CONFLICT providing Grant for University India State, acknowledges Telangana S.M. Zaheerabad, (TOIL), seeds. Limited Pongamia India Oils Tree acknowledge gratefully We Na to expression leading gene ultimately ACKNOWLEDGMENTS transporter effect stress, regions. downstream and apoplasmic salinity (antioxidant) The the defense under stress. in the salinity homeostasis sequestration hand, and influence under redox other nutrients could homeostasis of conferring the pathways ionic supply On patterns signaling constant and stress. provides above salinity osmotic polyols the high and conferring of to acids in plants amino involved Pongamia sugars, SA also a of JAs, are as adaptation phytohormones such three the metabolites that in in of hypothesize role induction we mechanisms crucial results, play tolerance correlation ABA our salt and on on Based signaling. focusing JAs including explicitly study Ca first induced wall in the cell role is 2013). of possible al., This permeability et its and (Zhao selectivity suggests environment the expression saline enhance high gene to CONCLUSIONS under wall CAD1-like transport cell the water the and in of solute increase rigidity for and consistent strength The the 2007). reinforcing al., et (Lee 2iae eepoe rmfiebooia elctsi oto n attetdpat.Errbrrepresents bar arrows Error Red NaCl plants. respectively. mM 8DAS salt-treated 300 and and control of 4 in leaves 1, (P replicates NaCl of biological mM sections five mean 500 cross from leaves the F), pooled of (A, and were sections color). E images cross Na (red 12 (D, I) measure iodide plants, and to propidium control H drawn and of (G, were leaves 8DAS, color) and of (green 4 sections AM 1, cross CoroNa-Green C) with and stained B were leaves of images S3. Figure < .5,* (P ** 0.05), ± 2+ + ffc fsl teso htsnhtcpromne hoohl ursec,clrpylcontent chlorophyll fluorescence, chlorophyll performance, photosynthetic on stress salt of Effect luiaino Na of Illumination ffc fsl teso ln morphology plant on stress salt of Effect D(=2.TowyAOAts a efre omauePvle s(o infiat * significant) (not ns P-values measure to performed was test ANOVA Two-way (n=12). SD /Cl eesntol oneatteNa the counteract only not levels + < < - ntelae n ot fcnrladsl-rae lns H n I aitosi the in variations (I) and (H) plants, salt-treated and control of roots and leaves the in .1 n (p * * * and 0.01) .5,* (P ** 0.05), n Na and ± + + /Ca D(=) w-a NV etwspromdt esr -ausn (not ns P-values measure to performed was test ANOVA Two-way (n=6). SD ursec nest.Qatfiaino Na of Quantification intensity. fluorescence < + .1 n (p * * * and 0.01) 2+ o ursec nlae fcnrladsalt-treated and control of leaves in fluorescence ion aisi h evsadroso oto n attetdpat.Error plants. salt-treated and control of roots and leaves the in ratios < .0)respectively. 0.001) 2+ o otn ntelae n ot fcnrladsalt-treated and control of roots and leaves the in content ion + < oiiybtas ciaenmru inln pathways signaling numerous activate also but toxicity .0)respectively. 0.001) 15 μ o m mol - o otn ntelae n roots and leaves the in content ion + ursec nest oethan more intensity fluorescence -2 ogmapinnata Pongamia s -1 ,()cagsi maximal in changes (B) ), .pinnata P. + o xlso and exclusion ion Confocal . .Salinity .. Posted on Authorea 27 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158800944.47703853 — This a preprint and has not been peer reviewed. Data may be preliminary. ieetslnt tescniin Cnrl,30ad50m ala ,4ad8A.Sxbiological Six 8DAS. and was test 4 ANOVA two-way 1, and at metabolite S3. each NaCl (P Table of * mM significant) changes (not 500 fold ns relative P-values and measure the to 300 measure performed biological (Control), to used Six 0 were 8DAS. conditions replicates and was stress test 4 salinity ANOVA two-way 1, different and at metabolite each NaCl S2. (P Table of * mM significant) changes (not 500 fold ns relative P-values and measure the to 300 measure performed respectively. (Control), to 8DAS used 0 and were conditions 4 replicates 1, stress at salinity NaCl different mM 500 and S1. 300 Table 0, reference conditions using stress by salinity normalized different gene each three of levels S11. of expression Figure The 8DAS. and 4 rRNA. 1, 18s at gene NaCl of mM roots 500 and and leaves 300 in expression genes enzyme plants S10. salt-treated Figure of (B) roots and mean (A) the leaves represents in CAD6, and of CAD2, PAL2, PAL1, POD, Cu/ZnSOD1-like, respectively. of 8DAS and S9 4 1, Figure at NaCl mM 500 rRNA. and 18s 300 gene 7) reference Figure using plementary by normalized gene S8. each Figure of correlation. levels of of degree expression indicates The scale 8DAS. color with of The represented bar. roots replicates and scale biological the row six (1, to horizontal of S7. points according single average time Figure blue a different an or three of is and red form value NaCl) codes the mM metabolite color in 500 Each different represented and respectively. (300 is 8DAS) treatments metabolite salt and correlation. two each 4 of of of columns degree 12) six indicates to with scale (1 portioned color with values The represented change bar. fold replicates relative scale biological the row six (1, to S6. horizontal of points according single average time Figure blue a different an or three of is and red form value NaCl) codes the mM metabolite color in 500 Each different represented and respectively. (300 is 8DAS) treatments metabolite salt and two each 4 of of columns 12) six to with (1 portioned values (P change mean fold * relative the significant) (not represents S5. ns bar P-values Figure Q Error to measure (P, drawn plants. to 8DAS, were salt-treated and performed arrows and 4 was White control 1, test respectively. (P ANOVA NaCl in 8DAS Two-way 4 mM and replicates images (n=12). 1, 300 magnified 4 biological SD NaCl L) of 1, five and mM roots NaCl K from 300 of mM (J, of pooled images 8DAS, 500 roots and magnified roots 4 of O) Na of 1, sections and measure NaCl images cross N mM magnified F), (M, B 500 R) and plants, roots (A, and of E control color). sections (D, (red of cross plants, iodide roots I) control propidium of and of and H color) roots (G, 8DAS, (green of and AM sections CoroNa-Green cross with C) stained and were roots of images S4. Figure < .pinnata P. .pinnata P. .1 n (p * * * and 0.01) eaiecnetainadfl hne fmjrmtbltsi evsof leaves in metabolites major of changes fold and concentration Relative eaiecnetainadfl hne fmjrmtbltsi ot of roots in metabolites major of changes fold and concentration Relative ito l 1mtbltso 0 MNC rae evsof leaves treated NaCl mM 300 of metabolites 71 all of List + ersnaiefiueo eiqatttv C nlsso rnpre ee xrsini leaves in expression genes transporter of analysis PCR semi-quantitative of figure Representative luiaino Na of Illumination eaiemN xrsinlvl fcl-alezmsadatoiatezmsi leaves in enzymes antioxidant and enzymes cell-wall of levels expression mRNA Relative . .pinnata P. h aditniiso rnpre ee fsm-uniaiePRaayi soe in (showed analysis PCR semi-quantitative of genes transporter of intensities band The etmpo eaoiecagsi evsof leaves in changes metabolite of map Heat etmpo eaoiecagsi ot of roots in changes metabolite of map Heat ersnaiefiueo eiqatttv C nlsso elwl nye n antioxidant and enzymes cell-wall of analysis PCR semi-quantitative of figure Representative ursec nest.Qatfiaino Na of Quantification intensity. fluorescence h aditniiso A1adCD-iegnsi evsadrosof roots and leaves in genes CAD1-like and CAD1 of intensities band The ne atSrs odtos Log Conditions. Stress Salt under t1 n DSrsetvl hncmae otercrepnigcnrl.Errbar Error controls. corresponding their to compared when respectively 8DAS and 4 1, at ± < D(n=6). SD .0)respectively. 0.001) ne he ieetslnt tescniin ,30ad50m ala ,4and 4 1, at NaCl mM 500 and 300 0, conditions stress salinity different three under nlae n ot of roots and leaves in + o ursec nroso oto n salt-treated and control of roots in fluorescence ion .pinnata P. .pinnata P. 2 odcagso A4 eO,MSD Cu/ZnSOD1, MnSOD, FeSOD, CAT4, of changes fold 16 < < .5,* (P ** 0.05), (P ** 0.05), + .pinnata P. ne he ieetslnt tescniin 0, conditions stress salinity different three under ne he ieetslnt tescniin 0, conditions stress salinity different three under .pinnata P. ursec nest oeta 2iae were images 12 than more intensity fluorescence < < .1 n (p * * * and 0.01) (p * * * and 0.01) ne atsrs.Log stress. salt under ne atsrs.Log stress. salt under .pinnata P. .pinnata P. .pinnata P. t1 n DSand 8DAS and 4 1, at .pinnata P. < < .0)respectively. 0.001) respectively. 0.001) .pinnata P. 2 2 aiso the of ratios aiso the of ratios ne three under ne three under < Confocal . .5,** 0.05), under Sup- ± Posted on Authorea 27 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158800944.47703853 — This a preprint and has not been peer reviewed. Data may be preliminary. di,S,Hsa,S . lKry .A,Kml .B . aa,S,Auiaa .A,&E-oyt,F. El-Domyati, & A., O. O., Abuzinadah N. S., Gadalla, Rabah, M., H., A. B. Shokry, K. A., Kamal, R. Younis, A., M., M. Al-Kordy, A. Alzohairy, M., S. impact A., mi- Hassan, Ramadan, fluxes S., M., green chloride Edris, S. Vacuolar in J. (2016). Sabir, stress. stress A. A., salinity Sirisattha, Bahieldin, De-Angeli, salinity early & & during under E., distribution Martinoia, H., dynamics and Na T., content Kageyama, Tohge, molecular of ion C., S., and role Eisenach, U., Sirisattha, The production Baetz, T., lipid (2017). Praneenararat, Enhanced W. H., M. croalga (2019). Iwahashi, Yaish, W. Y., & R. Syaputri, R., T., Al-Yahyai, Atikij, H., Saneoka, A., in Ueda, K oxidase V., polyphenol D. salinity chloroplasts with Assaha, isolated associated in process enzymes Physiological Copper (2017). (1949). Physiology Y. I. Wu, D. & W., Wang,Arnon, family. L., I. half-sib Y. Mott, Wang, alfalfa L., an D., M. in M. 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Quinn, gene Y., Guan, Yiu, Tao, tolerance F., Y., Y., salt T. Wang, K. novel Chan, A. C., Yim, a W., Communications Li, C., S. of C., Song, Tong, Identification Xu, G., X., (2014). Shao, S., Liu, M., H., K. Ni, T. Wong, X., Phang, F., Liu, Na (2016). M., C. M. Xie, Wong, X. W., S., Du, M. & Li, L., X., Y. Qi, Lu, F., ( W. cotton upland Gong, in E., 34548. tolerance Z. 6, stress Pan, salinity L., to J. related Sun, lization P., S. He, Z., Peng, ,17. 7, . giutrlResearch Agricultural N Research DNA 3 73-82. 13, ,4340. 5, urn pno nMicrobiology in Opinion Current odu vulgare Hordeum 2 471-483. 22, ,207-214. 2, + oslnt stress. salinity to ) APs ciiyi ottsusealsK enables tissues root in activity -ATPase ln inln Behavior & Signaling Plant LSOne PLoS ora fPatPhysiology Plant of Journal - cuuainin accumulation ln elReports Cell Plant 23 ,e38992. 7, ,208-210. 5, itcavera Pistacia ospu hirsutum Gossypium ln Physiology Plant .thaliana A. Plants 8 847-867. 38, ln Journal Plant ,e24259. 8, . rootstocks. L.) 6,1189-1202. 161, ,39. 8, 7,2445-2458. 172, otclsdrn atsrs is stress salt during cells root plants. ) + eeto n mediates and retention ieeg Research Bioenergy 3 131-142. 23, ln Environment Plant + cetfi Reports Scientific compartmenta- Pongamia Nature γ- 8, Posted on Authorea 27 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158800944.47703853 — This a preprint and has not been peer reviewed. Data may be preliminary. uea .(07.Asii cdadaitcsrs signaling. (2010). stress V. abiotic F. and ccid Breusegem, Abscisic & Chiwocha, (2007). Inz´e, UDP-glucosyltransferase D., I., N. A., Tuteja, the Clercq, K. De by Stubbs, B., homeostasis B., Cotte, Genty, acid de Arabidopsis W., van indole-3-butyric Boerjan, K., Vandenbroucke, of E., K., Perturbation Prinsen, Morreel, R., V., Fenske, O. S., Aken, B., V. Tognetti, re- messenger. by and Calcium-nutrient rootstocks (2019). almond K. of Thor, exchange. tolerance gas salt and enhances activity 98-105. jasmonate antioxidant 234-235, Methyl phytohormones, (2019). endogenous S. gulating Karimi, mechanisms & species, molecular tree V., Unravelling Tavallali, biofuel (2016). promising R. a R. in Attipalli, input biosynthesis analysis. & Auxin lipid scriptome T., (2014). to K. K. initiation Singha, floral B. M., from Zolman, in Shalini, & expression V., R. gene A., Sreeharsha, biosynthetic R. Rampey, auxin D., in changes J. by Cohen, Physiology P., mitigated Yu, are of A., disruptions roles pathway Hausman, and M., distribution Diversity, stress. G. (2015). abiotic Spiess, A. under Savoure, halophytes halophytic & in its T., accumulated Flowers, and A., compounds beet Bouchereau, osmoprotective carbohydrate C., of Abdell, case I., the Slama, shock salt vs. stress W., Salt Treder, W´ojcik, (2019). K., K., ancestor. J. Klamkowski, Tyburski, J., Kesy, & J., Niedojad A., Tretyn, species lo, K., tree Kurnik, key M., Go biofuel and lebiewski, M., potential metabolites Skorupa, a of pinnata, Dynamics Pongamia in (2019). of Products R. role seeds R. and their Attipalli, developing Crops of and & the S., developments in role Marriboina, current proteins The V., regulatory R. jasmonates: Sreeharsha, to (2015). T., K. response S. Singha, Plant Guo, (2019). & A. M., stress. Husen, environment. Zhong, salt changing & under Y., S., membranes Tang, K. thylakoid of Siddiqi, W., Reduced acids Zhang, fatty (2015). and J., H. ( proteins J. Sun, of rice Zhang, Reports J., regulation & upland the X., Chen, in in S. Y., exudation putrescine Cui, Yuan, ABA J., S., Y. by adaptation. Xia, Shu, caused drought R., is enhanced Liu, this system G., (2018). and Y. root B. 1) Liu, the Gaoshan Zheng, H., in & N. Ye, accumulation K., M., ABA A. M. Guo, Thukral, L., R., Shi, in Bhardwaj, detoxification K., insecticide M. stimulates acids Science Kanwar, fatty treatment Plant talk, H., observed cross Yuan, acid hormonal V., on Jasmonic acid Kumar, salicylic A., of safflower. Effects Sharma, salt-stressed (2019). from V. homeostasis Niknam, ions of & and application E., Foliar profile, H. (2015). Maboud, J. F., I. Shaki, Lee, & Biochemistry M., S. and in Kang, Physiology changes M., Plant physio-hormonal Hamayun, induced L., genotypes stress. jasmonate A. Maize salt methyl Khan, (2015). of M., F. phase Waqas, H. first R., the Bakhat, Shahzad, during & accumulation A., acid Science Fatima, jasmonic Crop F., in and M. vary Agronomy Qayyum, resistance H., salt Ali, in B., differing Pitann, N., A. Shahzad, ,14390. 5, M ln Biology Plant BMC 6,1092-1104. 165, rhtcueadwtrsrs tolerance. stress water and architecture ,1609. 9, cetfi Reports Scientific 4,111621. 140, ultno h ainlRsac Centre Research National the of Bulletin 0,443-451. 201, 9 57. 19, 6 406-416. 96, ,34315. 6, rnir nPatScience Plant in Frontiers ln Cell Plant ln&Cl Physiology Cell Plant& 24 iu sativum Pisum ora fPatInteractions Plant of Journal ln inln Behavior & Signaling Plant 2 2660-2679. 22, 3 153. 43, ne ies eprtr regimes. temperature diverse under rsiajuncea Brassica naso Botany of Annals 6 951-964. 56, ogmapinnata Pongamia 0 440. 10, ora fPatPhysiology Plant of Journal UGT74E2 rz sativa Oryza 4 340-346. 14, ,135-138. 2, Arabidopsis L. 1,433-447. 115, rnir in Frontiers Industrial . sn tran- using ora of Journal modulates Scientific .var. L. . Plant Posted on Authorea 27 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158800944.47703853 — This a preprint and has not been peer reviewed. Data may be preliminary. em .V,Zag . etrn,C 22) attlrnemcaim fplants. of mechanisms tolerance Salt (2020). T. C. in Nishio, Testerink, salinity & & Biology in Y., H., increase Zhang, Kitashiba, V., sudden M., E. to Nanzyo, Zelm, S., response Nasu, in K., roots Chow, and damage H., leaves oxidative napus B. of Brassica Kwan, from analysis P., plants transcriptome E. rice Comparative Kok, (2014). protects Z., stress. Zou, abiotic acid Y., and salicylic H. biotic Yong, Endogenous as (2004). well C. as responses. Mei, aging salt-stress by & caused plant M., stress mediating Qi, salt mechanisms Y., molecular regulate Yang, the negatively Elucidating B (2018). and Phytologist Y. A New Guo, Phytochrome & (2018). Y., D. Yang, metabolism Xi, Lignin & (2018). H., F. X. Lin, 2381-2393. Yang, J., & Li, J., of J. R., tolerance Yuan, Lv, M., T., H. Zeng, Yang, W., D. Qiu, B., ABNORMAL Y. involves (2017). in Liang, K. factors Yi, Y., rice. & environmental C. in Z., Yang, activity oxygen of Chen, reactive meristem J., roles proper root Luo, maintain The J., for to Peng, biosynthesis levels (2019). acid F., species Wang, salicylic Y. M., in Cai, Deng, functions two MERISTEM1 W., & INFLORESCENCE Ruan, on Q., H., 6-benzyladenine Zhao, Wang, L., exogenous Z., Xu, plant. of Tang, in Alleviation N., stress oxidative (2014). Chen, of D. Z., regulation He, Zha, X., & Xie, S., Yang, J., ( eggplant Chen, of J., genotypes He, X., Wu, L., Na Wang, (2018). N., Botany Bazihizina, S. Experimental S., Shabala, of Cai, & Q., Wu, Z., N., Chen, Su, Na C., S., Pandolfi, Mancuso, Y., M., Huang, E., Zhou, Azzarello, L., Shabala, Na H., and Wu, tolerance salt Plant (2018). (2016). H. M. Wu, H. Lam, soybean. & in B., accumulation Yu, chloride A., regulating Liu, L., Wang, Ca P., nonselective Wei, GMP-activated Lee, in H., cyclic genes Kollist, of I., CNGC6 Identification Puzorjova, and M., (2013). CNGC5 578-590. and Han, I. associated regulation N., J. and plant Robert, channels Schroeder, M., in cation & H. acid Ren, I., jasmonic N., Mori, of Nishimura, S., Functions S., (2020). Munemasa, F., M. Y. Li, Wang, & J., stress. Phyto- Xu, abiotic (2016). X., to Y. jasmonate Gong, Zhou, response acid-dependent Li., & abscisic J., Song, via Yu, J., K., tolerance Wang, Shi, cold X., regulate Xia, to J., antagonistically Zhou, methy- function E., signaling. its Onac, B and M., and Wang, acid A H., jasmonic chrome Li, by Z., growth Guo, plant F., Wang, cytokinin-induced of Inhibition (1982). jasmonates J. of lester. Kato, Effects roots. (2013). & sorghum U. J., S. in Park, Ueda, biosynthesis & sorgoleone C., for S. Chae, genes T., of Ecology W. expression Chemical Park, and B., accumulation Y. Kim, sorgoleone A., on A. Thwe, R., M. Uddin, + eusrto nroscne ieeta atsrs oeac ewe uu n ra wheat. bread and durum between tolerance stress salt differential confer roots in sequestration hsooi Plantarum Physiologia 1 24.1-24.31. 71, ortscinerea Botrytis ln Physiology Plant ioin tobacum Nicotiana yRNA-seq. by 1,523-539. 217, 9 712-722. 39, oau melongena Solanum c1idcddfnersos ntomato. in response defense BcG1-induced 9 3987-4001. 69, nentoa ora fMlclrSciences Molecular of Journal International 7,459-471. 170, iMdRsac International Research BioMed 4 249-252. 54, i B-amncai yegsi cross-talk. synergistic acid ABA-jasmonic via iMdRsac International Research BioMed + il)got ne atstress. salt under growth Mill.) rnir nPatScience Plant in Frontiers esn n transport. and sensing ln Cell Plant ln Journal Plant 25 GmCLC1 9 560-574. 29, + 04 467395. 2014, Arabidopsis xrso rmtectsladtissue-specific and cytosol the from extrusion 0 909-919. 40, ofr nacdsl oeac through tolerance salt enhanced confers M ln Biology Plant BMC h rpJournal Crop The 1 1446. 21, 09 9732325. 2019, 5 1082. 25, ur cells. guard Protoplasma ln elPhysiology Cell & Plant nulRve fPlant of Review Annual ln Physiology Plant ,215-225. 6, 8 103. 18, 5,169-176. 251, 2+ ora of Journal permeable Journal 163, 59, Posted on Authorea 27 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158800944.47703853 — This a preprint and has not been peer reviewed. Data may be preliminary. DadK B,( n )MJ,( n )S,( n )AAad( n )J nlae n ot of roots and respectively. leaves 8DAS in and JA O) 4 and points1, IAA, (H time J) and different and ABA three (P (C N) three at * zeatin, and at plants significant) mean (G I) treated (not NaCl SA, the and NaCl mM M) represents (B mM and 500 bar hormones 500 (F Error and endogenous and MeJA, of 300 L) 300 levels (control), and (control), The (E 0 0 IBA, 8DAS. concentrations K) and salt and 4 different (D 1, three points with time different treated days 30 after of roots 1. America Figure 1 of dehydrogenase States alcohol United cinnamyl the of LEGENDS of function G., FIGURE Sciences of M. of Loss Hahn, Academy (2013). in L.A., National defect A. Jackson, growth the R. temperature-sensitive C., of Dixon, a Fu, and & L., lignin Carpita, unconventional J., Gallego-Giraldo, in to A., Ralph, S., leads tolerance W. F., Pattathil, stress Tao, Chen, R., E., salt H., Zhou, Y. 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mM

A

GlcNAc Ba, CAA, Glu, ShiA Oxa Ala, HA, GlcNAc PA, LA, SA, Eta, IBua 5 Glu, HPip,

,

Eta,

GABA, PA, ,

Fru Oxopro

Rib, Hse pGlc oC , pTag

lI MytA Oxa , Oxopro

Phe Gri , , , Pdo lI

, StA

Ser

La, Cys

, Xyl Thr

,

, , Pro,

, NA,

Pdo ,

TA, GA, ,

, ,

,

CAA,

, Gri

, ,

Pi, pTag Pyr

, Bta Fa, Phe Grl 1DAS

Fru

, Ser Ara

MP,

,

Ara

Bta HA, , TA,

-

, StA

, pGlc g,

, 1DAS Hse , MytA , l ,

5

Asn , LA , GA,

Glta Gal,

HPip, Rib,

CA, MT, Man,

NA, ,

, pFru

,

Thy, ,

MP, , ThA

,

Suc

MT,

MI, MI, Met Thy,

Pi, GABA,

ThA IBua Glc 1DAS ,

,

G ,

Suc

- La CA,

Xyl Phytol, SA L PI, l Phytol,

3

,

- p,

pFru Asn , Asu ,

AA, SA, Glta 1DAS PI,

Asp, AA, ,

, ,

ShiA

Gal, Thr Ala, oC

, , ,

II l Group H

K

E lI

lI II l Group

B

Gro MytA PA, GA, Ala, Phe Oxopro Leu ThA Ala, AA, Phytol, Oxopro StA Gal, MT,

, ,

, Thy,

FA, GABA, Met , IBua

,

,

G Met Oxa NA, La,

pGlc MytA pTag ,

3 -

CAA, Ba,

Rib, p, Gal, ,

, HA, -

, LA,

Gal, Pip,

Bta

Asp, GABA, , ShiA

, , Glta

29

HA,

Pyr

Gal, BA,

Glc Oxa Hse Asu pFru ,

4DAS Fa, Rib,

Thy, ,

, , 5 ,

, Pro Pyr l pGlc , Gri HPip,

,

CA,

Thr Ara, Ba, 4DAS , LA, Pdo oC

Gro, l

PI,

oC , ,

Thr

,

, MT

ThA

, Met

Grl CA, ,

Asn ,

MA,

Eta

Phytol, GA, AA, GlcNAc , 5 G

HPip, - Leu - , 3 g,

SA,

,

Cys p, 4DAS

Suc Gri

Pdo

Suc Fa, PI,

,

,

BA, ,

Hse , pFru

4DAS

Man,

, Asn

, Xyl Asu ,

Ara, Fru Eta,

SA, FA, Pro,

, SA, ,

,

, , ,

Glta CAA,

MI Phe

Glu, PA,

Pi,

,, ,

II l Group

L I Group II l

F

C lI

Ba, Grl Glu, G Pyr MT, Phytol, PI, lI

3

p, -

,

g, SA,

Pi,

GlcNAc

Oxopro Asu

MP, pFru Oxopro Bta MytA La, Phytol, Suc Phe HA,

MI, AA, La,

, , Fru

,

, Rib,

, , Fru

Oxa Pi,

CAA,

Asp, LA, Ba, BA, , Eta, ,

Val, ,

, Ser

Gri

Asp,

Grl , Pdo ThA

,

Hse

Met

Ara, MP, Man, Pdo

, Glta

-

8DAS Bta g, Thr Ser Gro

, , , Fa,

l

-

oC

GlcNAc Gal, PI, , , G 8DAS

,

Pyr Met ,

,

pGlc , Rib,

Ery

, 3 Asn MT, Thr

l 5 SA, p, pGlc Gro,

, HPip, -

Ala, ,

SA,

MI, , ,

PA MA, ,

HA,

pTag Val, Gal, Eta, ,

5

,

Leu Met

HPip, Man, Met

MytA

ThA

Gri

Asu 8DAS Oxa Thy, Glta 8DAS , Thy, , -

- SA, Glc Gal, Gal, , ,

pTag ,

pFru , IBua

Leu ,

BA, NA,

Phe

Pro, Glu,

GA, PA,

Fa, , , , ,

,

Posted on Authorea 27 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158800944.47703853 — This a preprint and has not been peer reviewed. Data may be preliminary.

IBA Root J

IBA G Leaf IBA IBA

D A

JA JA

JA Pdo JA

Pdo Pdo Pdo

5 00

5 1DAS

3 00 1DAS 300 00 1DAS 1DAS mM mM mM mM

Phe Glu Phe Glu

Phe Glu Glu Phe

K

H

IBA IBA IBA

IBA B E

JA 30 JA

JA Pdo JA

Pdo

Pdo Pdo

5 00 4DAS

3

5 00 4DAS mM 00 4DAS mM 300 4 mM

DAS

mM

Phe Glu Glu Phe

Phe Glu Glu Phe

L

IBA IBA I

IBA IBA F C

JA JA JA

JA

Pdo Pdo

Pdo Positive correlation

N Pdo

3 egative correlation 00 8DAS

300

5 8DAS 00 mM 8DAS

5 mM 00 8DAS mM

mM

N Positive correlation

egative correlation

Phe Glu Oxopro Phe Glu

Phe Glu Phe

Glu

Posted on Authorea 27 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158800944.47703853 — This a preprint and has not been peer reviewed. Data may be preliminary.

SOS pathway stress Salt Ca -6.0 balance A Ionic 2+

300mM 1DAS 300mM 4DAS 300mM 8DAS 300mM 1DAS 500mM 4DAS 500mM 8DAS 500mM

Ion transporterIon

Leaf -4.0

gene

-2.0 0.0 JAs Antioxidantgenes 2.0 (SOD/CAT/POD) SALINITY TOLERANCE homeostasis

Redox 4.0 PM CNGC17 CNGC5 PM V PM CDPK3 CDPK32 V CCX1 CLC1 SOS3 V HKT1:1 SOS2 - NHX1 SA - H - H ABA CHX1 - - - + H H H +

31 6.0 ATPaseB subunit

+ ATPaseB subunit + +

ATPase 4.1 ATPase 4.1 ATPase 1

8.0

B -

like

Root Root growth and apoplasmic

sequestration

-2.0

H2O2 mediated cell wall lignification -1.0

Na 0.0 +

1.0 Sugars,sugar alcohols, amino acids, organic acids, fatty acids, fatty acids 2.0 Osmotic Osmotic balance PM PM V CDPK32 PM V CNGC17 CDPK3 V SOS3 SOS2 CLC1 HKT1:1 NHX1 - - - H 3.0 H CHX1 - - - + + H H H

CNGC5

ATPaseB subunit ATPaseB subunit + + +

ATPase 4.1 ATPase 4.1 ATPase 1 CCX1

4.0

-

like