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[(mechanisms)-332(allow)-336.4(EVs)-329.7(to)-331.9(ef)]TJ /F14 1 Tf 11.4646 0 TD ()Tj /F10 1 Tf 0.5291 0 TD [(ciently)-336.5(transport)-331.2(sensitive)-335.5(regulatory)]TJ -11.9937 -1.3921 TD [(molecules)-213.4(across)-206.9(extracellular)-218.6(environments)-210.9(and)-209.1(into)-211.2(recipient)-208.9(cells.)]TJ 0 -1.3858 TD [(In)-354.3(plants,)-353.8(however,)-350.7(the)-351(rigid)-355.3(nature)-355.4(of)-349.2(cell)-356.2(wall)-351.3(creates)-352.7(a)-350.4(physical)]TJ 0 -1.3921 TD 0.0243 Tc [(barrie)5.7(r)-525.1(to)-521.4(EV)-525.9(release.)-526.2(Eviden)4.7(ce)]TJ 14.5008 0 TD [(suggests)-518.7(that)-524(loc)5.8(a)-4.7(l)-519.3(cell)-525.6(w)2.2(all)]TJ -14.5008 -1.3858 TD 0.0329 Tc [(remodeling)-529.6(or)-529.1(loosening)]TJ /F11 1 Tf 10.9795 0 TD 0 Tc ()Tj /F10 1 Tf 0.9449 0 TD 0.0328 Tc [(potentiall)5.6(y)-536(t)0.9(riggered)-537(b)3.7(y)-529.7(s)-0.1(tress)-529.2(o)3.7(r)]TJ -11.9244 -1.3858 TD 0 Tc [(developmental)-467.5(cues)]TJ /F11 1 Tf 8.0252 0 TD ()Tj /F10 1 Tf 0.9197 0 TD [(may)-459.9(facilitate)-471.5(the)-458.1(passage)-468.5(of)-462.5(EVs)-468.3(into)-463.1(the)]TJ -8.9449 -1.3921 TD [(apoplast.)-395.3(Proteomic)-390.8(analyses)-392.8(have)-394.2(shown)-391.7(that)-393.3(plant)-387.8(EVs)-392.7(contain)]TJ 0 -1.3858 TD [(cell)-362.5(wall-modifying)-369.7(enzymes,)-365.3(such)-367.4(as)-364.3(glucanases)-363.9(and)-366.6(peroxidases,)]TJ 0 -1.3921 TD [(indicating)-231.7(that)-223.3(EVs)-229(may)-226.8(contribute)-231.7(to)-224.8(loosening)-226.7(the)-231.3(cell)-223.9(wall)-225.4(during)]TJ 0 -1.3858 TD [(secretion)-301.3(and)-303.6(transit)-302.4(\()]TJ 0.30196 0.72157 0.33333 scn 8.9008 0 TD [(Stotz)-298.6(et)-302.5(al.,)-300.6(2022)]TJ 0.15686 0.15294 0.16078 scn 6.652 0 TD (\).)Tj -13.9969 -1.3921 TD [(B)-9.1(u)0(i)-13.7(l)0(d)-11.1(i)0(n)-11.6(g)-512.1(o)-10.2(n)-511(t)-6.7(h)-10.7(is)-12.2(,)-512.7(h)0(e)-10.5(r)-7.7(e)-516.3(w)0(e)-519.6(e)0(x)-10.5(p)-9.1(l)-8.3(o)0(r)-11.6(e)-516.3(t)-6.7(h)0(e)-520.7(p)0(o)-13.1(t)-6.7(en)-13.1(t)-6.7(i)0(a)-14.8(l)-512.2(of)-519.2(EV)-13.4(s)-511.6(t)-6.7(o)]TJ -1.5559 -1.3858 TD 0.0251 Tc [(transport)-517(s)-1.5(peci)]TJ /F14 1 Tf 6.5386 0 TD 0 Tc ()Tj /F10 1 Tf 0.5543 0 TD 0.0237 Tc [(c)-524.1(r)-2.9(egu)-4.3(l)2.8(atory)-526.2(m)-2.1(olecu)-4.3(les)]TJ /F11 1 Tf 9.9654 0 TD 0 Tc ()Tj /F10 1 Tf 0.9386 0 TD 0.0244 Tc [(such)-528.1(as)-525(s)4.1(R)-4.1(NAs)-525(a)1.6(nd)]TJ -17.9969 -1.3921 TD 0 Tc [(sec)-13.4(onda)-11.4(r)-7.7(y)-505.8(m)-6.9(eta)-10.3(bo)-7.9(lit)-6.9(es)]TJ /F11 1 Tf 9.1528 0 TD ()Tj /F10 1 Tf 9 0 0 9 140.2582 168.8882 Tm [(t)-6.7(h)0(a)-8.4(t)-504.3(m)-6.9(odu)-9.6(lat)-12.6(e)-503.7(m)-6.9(icr)-12(obia)-10.1(l)-505.9(i)0(n)-11.6(ter)-7.8(ac)-10(t)-6.7(ion)-9.2(s)-505.3(a)0(t)]TJ -10.0724 -1.3858 TD [(the)-464.4(plant)]TJ /F11 1 Tf 3.7669 0 TD ()Tj /F10 1 Tf 0.5102 0 TD [(soil)-459.1(interface.)-471.2(We)-461.7(speci)]TJ /F14 1 Tf 9.789 0 TD ()Tj /F10 1 Tf 0.5354 0 TD [(cally)-463(highlight)-463.1(their)-464(potential)]TJ -14.6016 -1.3921 TD [(role)-466.8(in)-471.4(enhancing)-476.3(microbial)-465.5(nutrient)-476.8(mobilization)-467.5(and,)-476.1(in)-465.1(turn,)]TJ 0 -1.3858 TD [(improving)-305.7(plant)-299.6(nutrient)-306.7(uptake)-301.8(and)-303.6(nutrient)-300.4(use)-306.3(ef)]TJ /F14 1 Tf 20.9638 0 TD ()Tj /F10 1 Tf 0.5291 0 TD (ciency.)Tj /F1 1 Tf 13.5 0 0 13.5 304.8944 749.3668 Tm 0.30196 0.72157 0.33333 scn [(The)-300.4(role)-298.7(of)-300.2(the)-302.6(rhizosphere)]TJ 0 -1.1465 TD [(microbiome)-300.3(in)-301.9(the)-302.6(nutrient)-302.9(use)]TJ T* (ef)Tj /F2 1 Tf 0.8189 0 TD ()Tj /F1 1 Tf 0.5837 0 TD [(ciency)-305.4(of)-300.2(plants)]TJ /F10 1 Tf 9 0 0 9 318.8409 693.4109 Tm 0.15686 0.15294 0.16078 scn [(The)-316.2(rhizosphere)-325.8(microbiome)-312.7(critically)-322.4(regulates)-315.5(plant)-318.5(nutrient)]TJ -1.5496 -1.3921 TD 0.0469 Tc [(us)-11.2(e)-557.6(e)-3.2(f)]TJ /F14 1 Tf 2.7591 0 TD 0 Tc ()Tj /F10 1 Tf 0.5795 0 TD 0.0541 Tc [(ci)5.4(en)3(c)4(y)-552.5(\(N)5.7(UE\))-547.8(t)-3(h)5.6(r)2.3(ough)-548.8(proc)4(esse)4(s)-552(s)2.3(uch)-548.8(a)6.1(s)-552(n)3(utri)5.4(en)3(t)]TJ -3.3386 -1.3858 TD 0.0556 Tc [(mi)-5.7(neralizatio)-5(n)4.5(,)-557.9(s)-2.4(olubi)-5.7(l)3.2(iza)-5(t)4.8(i)-5.7(o)1.3(n,)-551.6(transpo)-5(r)3.8(t)-555.8(m)-1.7(od)-3.9(ulation,)-557.9(and)]TJ 0 -1.3921 TD 0.0135 Tc [(horm)-6(onal)-511.3(sig)5.4(n)-6.1(alin)-6.1(g)5.4(.)-518.1(Whil)5.2(e)-515.4(n)0.2(itrogen)-510.1(\()-2.6(N\))]TJ /F14 1 Tf 17.9906 0 TD 0 Tc ()Tj /F10 1 Tf 0.5417 0 TD 0.0155 Tc [(xa)5.3(tio)5.3(n)-514.4(pl)7.2(ays)-508.7(a)-511.3(role)]TJ -18.5323 -1.3858 TD 0 Tc [(primarily)-445(in)-446.2(legumes,)-451.5(a)-444.9(broader)-445.4(range)-448.6(of)-443.6(mechanisms)-451.7(involving)]TJ 0 -1.3921 TD [(microbial)-402.5(taxa,)-399.4(such)-398.9(as)]TJ /F15 1 Tf 9.8961 0 TD (Bacillus)Tj /F10 1 Tf 3.0551 0 TD (,)Tj /F15 1 Tf 0.6173 0 TD (Pseudomonas)Tj /F10 1 Tf 5.3858 0 TD (,)Tj /F15 1 Tf 0.6173 0 TD (Streptomyces)Tj /F10 1 Tf 5.0961 0 TD [(,)-399.3(and)]TJ -24.6677 -1.3858 TD [(arbuscular)-413.8(mycorrhizal)-414(fungi)-411.9(\(AMF\))-415.7(like)]TJ /F15 1 Tf 17.3102 0 TD [(Rhizophagus)-414.9(irregularis)]TJ /F10 1 Tf -17.3102 -1.3921 TD [(are)-288.5(more)-295.4(universally)-299.3(important)-295.8(in)-295.1(enhancing)-293.6(NUE)-294.4(\()]TJ 0.30196 0.72157 0.33333 scn 21.0457 0 TD [(Kuzyakov)-295.3(and)]TJ -21.0457 -1.3858 TD [(Razavi,)-303.3(2019)]TJ 0.15686 0.15294 0.16078 scn 5.0583 0 TD (\).)Tj -3.5087 -1.3921 TD [(Ni)-8.9(trogen)-505.6(miner)-13.3(alizatio)-10.4(n)]TJ /F11 1 Tf 9.915 0 TD ()Tj /F10 1 Tf 0.9197 0 TD [(mediate)-11.8(d)-500.5(b)0(y)-503.4(sapr)-9.5(otrophi)-8.8(c)-503.7(bacter)-9.1(ia)]TJ -12.3843 -1.3858 TD [(and)-354(fungi)-361.5(such)-361.1(as)]TJ /F15 1 Tf 7.6346 0 TD (Streptomyces)Tj /F10 1 Tf 5.4488 0 TD [(spp.)-362.2(and)]TJ /F15 1 Tf 3.8425 0 TD [(Trichoderma)-365(harzianum)]TJ /F11 1 Tf -16.926 -1.3921 TD ()Tj /F10 1 Tf 0.9134 0 TD [(involves)-349.8(the)-344.7(enzymatic)-345.1(degradation)-353.6(of)-342.8(organic)-350(matter,)-349(releasing)]TJ -0.9134 -1.3858 TD [(ammonium)-357.9(\(NH)]TJ 6.7499 0 0 5.9992 366.9731 547.37 Tm (4+)Tj 9 0 0 9 374.0598 543.4015 Tm [(\))-362.5(through)-359.3(protease)-359.7(and)-360.3(urease)-361.7(activity)-363.7(\()]TJ 0.30196 0.72157 0.33333 scn 16.2898 0 TD (Lazicki)Tj -23.9748 -1.3921 TD [(et)-365.5(al.,)-376.2(2020)]TJ 0.15686 0.15294 0.16078 scn 4.4976 0 TD [(\).)-371.3(This)-372.7(process)-375.7(governs)-369.8(the)-376.2(rate)-370.8(of)-368(N)-369.7(turnover)-377.5(and)-372.9(its)]TJ -4.4976 -1.3858 TD [(synchrony)-299.4(with)-296.1(plant)-299.6(demand)-300.4(for)-293.8(nutrients.)-304.2(In)-297.7(parallel,)-297.3(ammonia-)]TJ 0 -1.3921 TD 0.0612 Tc [(oxidizing)-558(b)0.6(a)-5.7(c)4.8(teri)-6.4(a)-560.1(\()1(e.g.,)]TJ /F15 1 Tf 11.7858 0 TD 0.0619 Tc (Nitrosospira)Tj /F10 1 Tf 5.5748 0 TD 0.6274 Tc [(\)a)566.8(n)570(d)0.9(a)566.8(r)563(c)571(h)566.3(a)560.5(e)571(a\()567.3(e)564.7(.)568.2(g)562.6(.)568.2(,)]TJ /F15 1 Tf -17.3606 -1.3858 TD 0 Tc (Nitrososphaera)Tj /F10 1 Tf 5.9654 0 TD [(\))-494.8(mediate)-496.8(nitri)]TJ /F14 1 Tf 6.2236 0 TD ()Tj /F10 1 Tf 0.5354 0 TD [(cation,)-494.5(in)]TJ /F14 1 Tf 3.9685 0 TD ()Tj /F10 1 Tf 0.5354 0 TD [(uencing)-495.2(nitrate)-496.1(\(NO)]TJ 6.7499 0 0 5.9992 537.2219 491.8676 Tm (3)Tj 0.4788 0.9166 TD (-)Tj 9 0 0 9 542.8345 493.3983 Tm (\))Tj -26.4378 -1.3921 TD [(availability)-302(and)-297.3(root)-305.6(uptake)-301.8(dynamics)-305.4(\()]TJ 0.30196 0.72157 0.33333 scn 15.8173 0 TD [(Prosser)-303.8(and)-303.6(Nicol,)-300.4(2012)]TJ 0.15686 0.15294 0.16078 scn 9.6441 0 TD (\).)Tj -23.9118 -1.3858 TD 0.0408 Tc [(Ph)4.9(osph)4.9(orus)-533.8(\(P\))-535.9(a)-0.9(va)5.4(ila)5.4(b)-0.9(i)4.7(l)1(ity)-534.3(i)-1.6(s)-533.8(e)-3(nh)4.9(anced)-535.3(b)-0.9(y)-534.3(p)0.2(ho)5.4(spha)5.4(te-)]TJ -1.5496 -1.3921 TD 0 Tc [(solubilizing)-359.5(microorganisms)-363.7(\(PSMs\),)-363.4(notably)]TJ /F15 1 Tf 18.6457 0 TD [(Bacillus)-359.8(megaterium)]TJ /F10 1 Tf -18.6457 -1.3858 TD (and)Tj /F15 1 Tf 1.8331 0 TD (Pseudomonas)Tj /F16 1 Tf 5.726 0 TD ()Tj /F15 1 Tf 0.5228 0 TD (uorescens)Tj /F10 1 Tf 3.7291 0 TD [(,)-336.3(which)-337.7(secrete)-342.2(organic)-343.7(acids)-340(such)-342.2(as)]TJ -11.811 -1.3921 TD [(gluconic)-349.2(and)-354(citric)-350.5(acid)-357.5(that)-349.2(chelate)-350.1(cations)-354.8(and)-354(release)-354.3(inorganic)]TJ 0 -1.3858 TD [(phosphate)-338.8(from)-332.2(mineral)-340.4(complexes)-341.9(\()]TJ 0.30196 0.72157 0.33333 scn 15.0299 0 TD (Rodr)Tj /F12 1 Tf 1.9969 0 TD ()Tj /F13 1 Tf 0.5543 0.0441 TD ()Tj /F10 1 Tf -0.0315 -0.0441 TD [(guez)-338(and)-335.1(Fraga,)-337.6(1999)]TJ 0.15686 0.15294 0.16078 scn 8.674 0 TD (\).)Tj -26.2236 -1.3858 TD [(AMF)-383.6(such)-392.6(as)]TJ /F15 1 Tf 5.8835 0 TD [(Rhizophagus)-396(irregularis)]TJ /F10 1 Tf 9.8394 0 TD [(form)-382.6(extensive)-390.5(extraradical)]TJ -15.7228 -1.3921 TD [(hyphae)-199.9(that)-204.4(acquire)-201.4(orthophosphate)-207.9(beyond)-202(the)-206.1(root)-198.6(depletion)-205.7(zone)]TJ 0 -1.3858 TD [(and)-202.8(transfer)-215.5(it)-206.5(to)-205.9(the)-206.1(host)-211.6(via)-205.6(the)-206.1(mycorrhizal)-212.4(phosphate)-212.8(transporter)]TJ 0 -1.3921 TD [(pathway)-467.9(\()]TJ 0.30196 0.72157 0.33333 scn 4.1764 0 TD [(Smith)-474.2(and)-467.3(Smith,)-476.7(2011)]TJ 0.15686 0.15294 0.16078 scn 9.7953 0 TD [(\).)-472.1(While)]TJ /F15 1 Tf 3.9307 0 TD [(R.)-475.6(irregularis)]TJ /F10 1 Tf 5.8079 0 TD [(is)-472.1(most)]TJ -23.7102 -1.3858 TD [(extensively)-413.2(studied,)-413.8(other)-413(species)-412.5(such)-411.5(as)]TJ /F15 1 Tf 17.4488 0 TD [(Funneliformis)-418.2(mosseae)]TJ /F10 1 Tf 9.115 0 TD (,)Tj /F15 1 Tf -26.5638 -1.3921 TD [(Claroideoglomus)-229.7(etunicat)]TJ /F10 1 Tf 10.0095 0 TD [(um,)-226.4(and)]TJ /F15 1 Tf 3.4961 0 TD [(Glomus)-227.7(versiforme)]TJ /F10 1 Tf 7.5654 0 TD [(have)-224.1(also)-219.1(been)]TJ -21.0709 -1.3858 TD [(shown)-209(to)-205.9(enhance)-216.9(nutrient)-212.2(uptake)-213.6(and)-209.1(soil)-207.1(stability,)-214.3(sometimes)-213.8(with)]TJ 0 -1.3921 TD [(crop-)-296.4(or)-301.3(environment-speci)]TJ /F14 1 Tf 11.1055 0 TD ()Tj /F10 1 Tf 0.5354 0 TD [(c)-295.8(advantages)-310.3(\()]TJ 0.30196 0.72157 0.33333 scn 5.7323 0 TD [(Shukla)-303.4(et)-302.5(al.,)-300.6(2025)]TJ 0.15686 0.15294 0.16078 scn 7.3575 0 TD (\).)Tj -23.1811 -1.3858 TD 0.0169 Tc [(M)-5.2(i)-0.3(crobial)-514.2(modu)-4.8(lation)-519.3(of)-517.3(root)-518.9(architecture)-518.3(is)-513.6(another)-519.9(k)2(ey)]TJ -1.5496 -1.3921 TD 0.035 Tc [(mechani)5.2(s)-4.2(m.)-534.4(Plant)-532.3(g)1.7(rowth-prom)]TJ 14.2551 0 TD 0.0357 Tc [(oting)-533.1(r)-3.4(h)6.1(i)-0.4(zobacteri)5.9(a)-535.2(\()0.7(PGPR\),)]TJ -14.2551 -1.3858 TD (including)Tj /F15 1 Tf 4.6173 0 TD [(Azospirillum)-531.3(brasilense)]TJ /F10 1 Tf 10.7213 0 TD 0.037 Tc (and)Tj /F15 1 Tf 2.1417 0 TD 0.0356 Tc [(Pseud)6.5(o)-0.3(monas)-532.2(p)-1.9(utida)]TJ /F10 1 Tf 9.0835 0 TD 0 Tc (,)Tj -26.5638 -1.3921 TD 0.0178 Tc [(synth)-5.5(e)-0.8(size)-523.7(aux)-5.5(i)0.6(ns)-519(\(notably)-519.5(i)0.6(ndole-3-acetic)-517.4(a)-5(c)-0.8(id\))-521.1(t)-1.5(hat)-518(i)0.6(ncrease)]TJ 0 -1.3858 TD 0 Tc [(lateral)-303(root)-299.3(proliferation)-311.5(and)-303.6(root)-305.6(hair)-304.1(density,)-303.7(expanding)-311.4(the)-300.6(root)]TJ 0 -1.3921 TD [(surface)-228.2(area)-223.2(for)-230.8(nutrient)-224.8(uptake)-232.5(\()]TJ 0.30196 0.72157 0.33333 scn 13.0268 0 TD [(Spaepen)-234.7(et)-226.9(al.,)-225(2007)]TJ 0.15686 0.15294 0.16078 scn 7.7606 0 TD [(\).)-226.4(Additionally,)]TJ -20.7874 -1.3858 TD 0.0325 Tc [(AC)-4.4(C)-539.8(d)-1.8(e)-5(a)-2.9(m)-5.9(i)-3.6(nas)-6.7(e)]TJ /F11 1 Tf 7.1685 0 TD 0 Tc ()Tj /F10 1 Tf 0.5417 0 TD 0.0359 Tc [(p)-4.7(r)3(odu)-4.7(c)4.7(ing)-532.9(m)-2.5(icrobes)-538.7(s)3.1(uch)-535.5(a)0.5(s)]TJ /F15 1 Tf 13.3354 0 TD 0.0356 Tc (Pseudomonas)Tj /F16 1 Tf -21.0457 -1.3921 TD 0 Tc ()Tj /F15 1 Tf 0.5228 0 TD [(uo)-15.2(re)-9.5(s)-7.2(c)0(e)-11.1(n)0(s)]TJ /F10 1 Tf 4.2835 0 TD [(l)-8.3(o)0(w)-7.2(e)0(r)-517.6(e)0(t)-12.7(h)0(y)-12.6(l)0(e)-8(n)-7(e)-516.3(l)0(e)-8(v)-8.1(el)-8(s)-511.6(u)-9.1(n)-7(d)0(e)-8.9(r)-511.6(n)-7(u)-9.1(tr)-8(ie)-10.6(n)-7(t)-510.6(s)-7.7(t)-6.7(r)-7.7(ess)-15(,)-512.7(t)-6.7(he)-10.5(r)-7.7(e)0(b)-10(y)]TJ -4.8063 -1.3858 TD 0.0241 Tc [(pr)-8.8(om)-8(ot)-7.8(in)-8.1(g)-525.8(r)-8.7(oo)-5(t)-530.6(e)-0.8(lo)-5(nga)-5(t)-1.5(i)-5.7(o)-5(n)-531(a)1.3(n)-8.1(d)-526.8(n)-8.1(u)-3.9(tri)-5.7(e)-0.8(n)-8.1(t)-530.6(-f)-6.2(or)-8.8(agi)-5.7(n)-1.8(g)-532.1(e)-7.1(f)]TJ /F14 1 Tf 23.578 0 TD 0 Tc ()Tj /F10 1 Tf 0.5606 0 TD 0.0257 Tc 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7.7039 0 TD [(\).)-314.6(In)-310.2(the)-313.2(context)-311.7(of)-311.4(nitrogen)-315.6(cycling,)-315.3(EVs)]TJ -10.148 -1.3921 TD [(could)-275.9(act)-274.9(as)-276.2(messengers)-285(that)-273.7(regulate)-282.6(microbial)-276.6(activity,)-277.9(potentially)]TJ 0 -1.3858 TD (in)Tj /F14 1 Tf 0.8 0 TD ()Tj /F10 1 Tf 0.5291 0 TD [(uencing)-432.2(the)-420.3(abundance)-434.2(and)-423.3(activity)-426.7(of)-431(nitrogen-transforming)]TJ -1.3291 -1.3921 TD [(bacteria.)-473.6(However,)-480.7(no)-470.8(studies)-479.2(have)-476.1(yet)-474.4(quanti)]TJ /F14 1 Tf 19.3953 0 TD ()Tj /F10 1 Tf 0.5291 0 TD [(ed)-475(the)-477(extent)-477.8(to)]TJ -19.9244 -1.3858 TD [(which)-331.4(EVs)-329.7(contribute)-338.8(to)-331.9(nutrient)-331.9(cycling,)-340.5(making)-335.3(this)-332(an)-332.2(exciting)]TJ 0 -1.3921 TD [(avenue)-298.6(for)-300.1(future)-308.2(research.)]TJ /F1 1 Tf 13.5 0 0 13.5 304.8944 640.233 Tm 0.30196 0.72157 0.33333 scn [(The)-300.4(role)-298.7(of)-300.2(extracellular)-305.5(vesicles)-304.5(for)]TJ 0 -1.1507 TD [(NUE)-299.1(of)-300.2(crops)]TJ /F10 1 Tf 9 0 0 9 318.8409 599.6975 Tm 0.15686 0.15294 0.16078 scn 0.0598 Tc [(Plant)-557.9(E)-1.8(V)-559.7(p)0.3(roteomic)-557.3(studies)-558.9(have)-557.3(detected)-560.4(numerous)]TJ -1.5496 -1.3858 TD 0.0297 Tc [(transporters)-526(\(e.g.,)-533.4(f)5.7(o)-5.7(r)-526(N)0.2(O)]TJ 6.7499 0 0 5.9992 409.3794 585.751 Tm 0 Tc (3)Tj 0.5207 0.9167 TD (-)Tj 9 0 0 9 415.559 587.2251 Tm 0.0304 Tc [(,)-526.4(p)2.4(hosphate,)-526.4(a)1.3(nd)-526.8(NH)]TJ 6.7499 0 0 5.9992 502.0723 591.2503 Tm 0.0417 Tc (4+)Tj 9 0 0 9 509.7258 587.2251 Tm 0.0294 Tc [(\),)-527.4(p)-4.9(r)2.9(oton)]TJ -22.7591 -1.3921 TD 0 Tc [(ATPases,)-350.6(and)-354(aquaporins)-353.3(in)-351.8(root-secreted)-359.6(EV)-347.3(samples)-358.2(\()]TJ 0.30196 0.72157 0.33333 scn 22.885 0 TD [(Cai)-354.1(et)-346.6(al.,)]TJ -22.885 -1.3858 TD (2018)Tj 0.15686 0.15294 0.16078 scn 1.915 0 TD [(\),)-409.1(supporting)-413.2(the)-407.7(idea)-414.2(that)-405.9(EVs)-411.6(may)-409.5(play)-413.7(a)-407.1(role)-410.1(in)-408.4(NUE)-407.8(of)]TJ -1.915 -1.3858 TD [(crops.)-288.8(Intensive)-293.4(studies)-296.5(have)-293.4(focused)-292.7(mainly)-290.8(for)-293.8(legumes)-291.5(and)-291(their)]TJ 0 -1.3921 TD [(respective)-395.5(symbionts.)-392.8(Root-exuded)]TJ /F14 1 Tf 14.5701 0 TD ()Tj /F10 1 Tf 0.5291 0 TD [(avonoids,)-397.2(delivered)-392.6(via)-394.6(ABC)]TJ -15.0992 -1.3858 TD [(transporters,)-474.7(initiate)-472.4(symbiosis)-470.7(by)-471.9(inducing)-471.4(bacterial)-473.1(nodulation)]TJ 0 -1.3921 TD [(genes)-299.5(\()]TJ 0.30196 0.72157 0.33333 scn 2.8409 0 TD [(Sugiyama)-301.1(et)-302.5(al.,)-300.6(2007)]TJ 0.15686 0.15294 0.16078 scn 8.5102 0 TD (\).)Tj -9.8016 -1.3858 TD [(A)-213.8(recent)-209.5(study)-210.8(showed)-210.9(that)-210.7(legume)-214.6(roots)-206.2(release)-215.7(EVs)-210.1(containing)]TJ -1.5496 -1.3921 TD [(small)-476(RNAs)-474.7(\(sRNAs\))-476.7(and)-479.9(peptides)-480.4(that)-475.2(may)-472.5(regulate)-477.9(nitrogen-)]TJ /F14 1 Tf 0 -1.3858 TD ()Tj /F10 1 Tf 0.5291 0 TD [(xing)-313.9(symbiosis)-319.5(\()]TJ 0.30196 0.72157 0.33333 scn 6.589 0 TD (Ayala-Garc)Tj /F12 1 Tf 4.5543 0 TD ()Tj /F13 1 Tf 0.2898 0.0378 TD ()Tj /F10 1 Tf -0.0315 -0.0378 TD [(a)-318.9(e)0(t)-315.1(al.,)-319.5(2025)]TJ 0.15686 0.15294 0.16078 scn 5.1465 0 TD [(\).)-314.6(Proteomic)-321.5(analyses)-317.2(of)]TJ -17.0772 -1.3921 TD 0.0198 Tc [(EVs)-523.3(f)-4.2(ro)-3(m)]TJ /F15 1 Tf 4.7811 0 TD 0.0205 Tc [(Lotus)-522.1(b)1.9(urtt)-5.1(ii)]TJ /F10 1 Tf 5.978 0 TD 0.0212 Tc (and)Tj /F15 1 Tf 2.085 0 TD 0.0203 Tc [(Phase)-6.7(o)3.3(lus)-522.3(v)-2.5(ulga)-7.2(ris)]TJ /F10 1 Tf 8.3339 0 TD 0.0211 Tc [(revealed)-523.5(that)]TJ -21.178 -1.3858 TD 0.012 Tc [(app)-3.4(r)-2(ox)-5(imately)-512.7(10%)-510.4(of)-515.9(EV-associa)-4.5(t)-1(ed)-513.7(protei)-5.2(ns)-512.2(are)-510.6(i)1.1(nvolved)-513.7(i)1.1(n)]TJ 0 -1.3921 TD 0.0104 Tc [(RNA)-505.7(m)-2.8(e)10.4(t)-8.6(abolism,)-508.6(inclu)-5(d)1.3(ing)-508(D)-3.9(EA)10.4(D)-9.8(/)10.4(D)-9.5(E)-0.8(A)10.4(H)-4(-box)-510.6(RNA)-512.1(h)10.4(e)-6.4(l)2.1(icases)]TJ 0 -1.3858 TD 0 Tc [(linked)-434.2(to)-438.9(sRNA)-436.9(processing.)-441.9(These)]TJ /F14 1 Tf 14.6331 0 TD ()Tj /F10 1 Tf 0.5291 0 TD [(ndings,)-442.9(along)-434.5(with)-441(evidence)]TJ -15.1622 -1.3921 TD [(of)-519.2(i)-10.9(n)-7(t)-6.7(e)0(r)-13.7(-)-11.4(ki)-13.2(n)-7(g)-8.1(d)-9.1(o)0(m)-521.1(R)-9.6(NA)-520.4(i)-10.9(n)-7(t)-6.7(e)0(r)-13.7(f)-11.4(er)-13.7(e)-12.3(n)-7(ce)]TJ /F11 1 Tf 15.1433 0 TD ()Tj /F10 1 Tf 0.926 0 TD 0.0092 Tc [(wh)9.2(ere)-507.1(p)0.1(lants)-508.7(u)9.2(se)-507.1(EVs)-502.4(t)2.5(o)]TJ -16.0693 -1.3858 TD 0.5242 Tc [(d)515.1(e)511.9(l)515.9(i)513.3(v)509.8(e)511.9(rs)516.5(R)508.3(N)513.6(A)512(s)0(t)517.5(o)-2.6(o)514(t)511.2(h)513.5(e)511.9(ro)514(r)516.5(g)509.8(a)514(n)510.9(i)513.3(s)510.2(m)511(s)]TJ /F11 1 Tf 14.8346 0 TD 0 Tc ()Tj /F10 1 Tf 0.9323 0 TD 0.5232 Tc [(s)515.5(u)507.8(g)515.1(g)508.8(e)510.9(s)515.5(tt)510.2(h)512.5(a)513(tE)512(V)508.4(-)511.8(m)510(e)510.9(d)514.1(i)512.3(a)512.9(t)510.2(e)510.8(d)]TJ -15.7669 -1.3921 TD 0 Tc [(sRNA)-506.2(transf)-12.8(er)-505(may)-510.3(modula)-9.8(te)-504(rhiz)-10.1(obial)-509.7(gene)-506(expr)-14.7(ession)-509.3(duri)-11.6(ng)]TJ 0 -1.3858 TD [(early)-298.6(symbiosis)-300.6(\()]TJ 0.30196 0.72157 0.33333 scn 6.7528 0 TD (Ayala-Garc)Tj /F12 1 Tf 4.5543 0 TD ()Tj /F13 1 Tf 0.2898 0.0378 TD ()Tj /F10 1 Tf -0.0315 -0.0378 TD [(a)-300(e)0(t)-302.5(al.,)-300.6(2025)]TJ 0.15686 0.15294 0.16078 scn 5.0961 0 TD (\).)Tj -15.1118 -1.3921 TD [(In)-310.2(addition)-313.5(to)-306.7(their)-312.8(role)-309.4(in)-314(nutrient)-306.7(signaling)-317.6(within)-307.7(the)-306.9(plant,)]TJ -1.5496 -1.3858 TD [(several)-249.3(miRNAs,)-249.3(including)-252.9(miR160,)-249.5(miR167,)-255.8(miR169,)-249.5(miR172,)-249.5(and)]TJ 0 -1.3921 TD 0.0164 Tc [(miR396,)-515.2(h)-0.6(ave)-518.8(b)-0.1(een)-519.8(s)2.4(hown)-513.5(to)-516.7(regulate)-512.5(nodule)-512.5(f)-1.3(ormation)-519.8(and)]TJ /F15 1 Tf 0 -1.3858 TD 0.027 Tc [(Rh)-4.7(iz)-4.2(ob)-4.2(ium)]TJ /F10 1 Tf 5.1087 0 TD 0.031 Tc [(i)7.5(n)-1.2(fe)6.1(ctio)8.2(n)-530.4(i)1.2(n)-524.1(l)3.8(egum)5.2(es)-524.7(\()]TJ 0.30196 0.72157 0.33333 scn 10.041 0 TD 0.0283 Tc [(Yousuf)-531.1(et)-532.7(al.,)-528.5(2021)]TJ 0.15686 0.15294 0.16078 scn 8.6299 0 TD 0.0292 Tc [(\).)-527.6(This)]TJ -23.7795 -1.3921 TD 0.0183 Tc [(undersc)-6.6(o)1.8(res)-518.5(t)-1(hei)-5.2(r)-518.5(dual)-519.1(ro)-4.5(le)-523.2(in)-517.9(b)-4.5(o)1.8(th)-515.3(nutrient)-517.5(a)-4.5(c)-0.3(qui)-5.2(s)-2(ition)-524.2(a)1.8(nd)]TJ 0 -1.3858 TD 0 Tc (plant)Tj /F11 1 Tf 2.0472 0 TD ()Tj /F10 1 Tf 0.5102 0 TD [(microbe)-461.1(communication.)-457.1(The)-454.7(packaging)-462.7(of)-456.2(such)-455.6(miRNAs)]TJ -2.5575 -1.3921 TD [(into)-356(EVs)-354.9(may)-359.1(represent)-360(a)-356.7(sophisticated)-363.9(mechanism)-362.1(to)-357(coordinate)]TJ 0 -1.3858 TD [(mi)-11.5(cr)-13.7(ob)-7.9(i)-10.9(a)0(l)-516.1(s)-7.7(y)-8.1(m)-6.9(bi)-8.5(o)-10.2(s)0(i)-12.2(s)-511.6(wi)-14.2(t)-6.7(h)-508.4(ho)-14.7(s)-7.7(t)-510.6(n)-7(u)0(t)-9.5(r)-7.7(i)-10.9(e)0(n)-13.1(t)-510.6(s)-7.7(t)-6.7(at)-10.6(u)-9.1(s)0(.)-514.1(I)-7.2(d)-9.1(e)0(n)-13.1(t)-6.7(if)-9.6(y)-8.1(i)0(n)-11.6(g)-512.1(E)-11.2(V)0(-)]TJ 0 -1.3921 TD [(associa)-12.5(ted)-506.9(sRNA)-8.6(s)-499(enh)-11.2(ancin)-9.7(g)-499.5(sym)-10.1(biotic)-508.4(ef)]TJ /F14 1 Tf 17.5118 0 TD ()Tj /F10 1 Tf 0.5354 0 TD [(cienc)-10.8(y)-499.5(coul)-8.5(d)-506.8(directl)-10.6(y)]TJ -18.0472 -1.3858 TD [(inform)-299.7(crop)-303.9(improvement)-306(strategies.)]TJ 1.5496 -1.3921 TD [(Although)-309.2(most)-302.3(mechanistic)-309.5(research)-307.7(has)-299.5(focused)-305.3(on)-307(symbiotic)]TJ -1.5496 -1.3858 TD [(systems,)-273.6(similar)-276.6(EV-mediated)-278.8(processes)-276(are)-275.9(predicted)-278.6(to)-275.2(operate)-276.5(in)]TJ 0 -1.3921 TD [(n)-7(o)0(n)-11(-)-11.4(l)-8.3(e)0(g)-14.2(u)-9.1(m)-6.9(e)-516.3(r)-7.7(h)-10.7(iz)-16.9(os)-11.6(p)-9.1(h)-10.7(er)-13.7(e)-12.3(s)-7.7(.)-512.7(I)-7.2(n)-517.3(t)-6.7(h)-10.7(e)0(s)-13.7(e)-516.3(s)-7.7(y)-8.1(s)-7.7(t)-13(em)-12.9(s)-7.7(,)-519(EV)-13.4(s)-517.9(d)-9.1(er)-13.7(i)-10.9(v)-8.1(ed)-525.4(fr)-12.7(o)-10.2(m)]TJ 0 -1.3858 TD [(rhizobacteria)-334(have)-337.5(been)-331.7(shown)-335(to)-331.9(encapsulate)-338.4(hydrolytic)-337.8(enzymes)]TJ /F11 1 Tf 0 -1.3921 TD ()Tj /F10 1 Tf 0.9134 0 TD [(such)-222.5(as)-225.8(phosphatases)-225.8(and)-221.7(proteases)]TJ /F11 1 Tf 14.0032 0 TD ()Tj /F10 1 Tf 0.9134 0 TD [(that)-223.3(catalyze)-225.4(mineralization)]TJ -15.8299 -1.3858 TD [(of)-292.5(organic)-293.3(phosphorus)-300.1(and)-297.3(nitrogen)-296.7(near)-295.5(root)-293(surfaces,)-301.3(potentially)]TJ 0 -1.3921 TD [(elevating)-241.5(localized)-238.9(pools)-240.8(of)-242.1(orthophosphate)-245.7(and)-240.6(NH)]TJ 6.7499 0 0 5.9992 491.1306 166.2236 Tm (4+)Tj 9 0 0 9 500.3715 162.1984 Tm [(in)-244.7(alignment)]TJ -21.7197 -1.3858 TD [(with)-321.3(root)-330.8(uptake)-327(zones)-333.1(\(microbial)-330.4(extracellular)-332(enzyme)-329.9(dynamics;)]TJ T* [(rhizosphere)-275.4(enzyme-turnover)-277.8(studies\))-274.7(\()]TJ 0.30196 0.72157 0.33333 scn 15.8173 0 TD [(Tian)-270.8(et)-271(al.,)-275.4(2020)]TJ 0.15686 0.15294 0.16078 scn 6.4252 0 TD [(\).)-270.5(EV)-271.7(cargo)]TJ -22.2425 -1.3921 TD [(may)-302.4(also)-307.3(include)-308.5(siderophores)-310.9(and)-303.6(redox-active)-311(molecules)-307.9(capable)]TJ 0 -1.3858 TD [(of)-324(chelating)-326(iron)-325.5(and)-322.5(mobilizing)-328.8(mineral-bound)-328.3(phosphorus,)-327.8(akin)]TJ 0 -1.3921 TD 0.0169 Tc [(to)-516.2(p)-4.8(h)-0.1(osp)-4.8(h)-0.1(ate-solubilizing)-520.4(microbial)-520.5(processes)-513.6(i)-6.6(denti)]TJ /F14 1 Tf 22.5827 0 TD 0 Tc ()Tj /F10 1 Tf 0.548 0 TD 0.0179 Tc [(ed)-520.4(for)-512.6(a)]TJ -23.1307 -1.3858 TD 0.0085 Tc [(nu)8.5(m)-7.5(b)8.5(er)-509.4(o)8.5(f)-510.7(st)-4.5(ra)8.5(i)-6.3(n)1.5(s)-503.1(f)-2.9(ro)8.5(m)-512.6(t)1.8(he)-501.5(gene)8.5(r)-5.2(a)]TJ /F15 1 Tf 15.6787 0 TD 0.009 Tc [(Bacil)9(l)-4.3(us)]TJ /F10 1 Tf 3.6157 0 TD 0.0086 Tc (and)Tj /F15 1 Tf 2.0284 0 TD 0 Tc [(P)-8.6(s)-7.2(e)-8.2(u)-10.7(do)-14.7(mo)-17.1(na)-13.1(s)]TJ /F10 1 Tf -21.3228 -1.3921 TD (\()Tj 0.30196 0.72157 0.33333 scn 0.3402 0 TD [(Suleimanova)-381.3(et)-371.8(al.,)-376.2(2023)]TJ 0.15686 0.15294 0.16078 scn 9.9906 0 TD [(\).)-371.3(Although)-378.4(direct)-380.2(miRNA)-377.7(regulation)-374(of)]TJ /F1 1 Tf 6.9999 0 0 6.9999 49.6063 793.6439 Tm 0.49804 0.49804 0.49412 scn [(Paponov)-303.2(et)-300.8(al.)]TJ 58.5567 0 TD (10.3389/fpls.2025.1681793)Tj -58.5567 -108.3259 TD [(Frontiers)-304.6(in)]TJ 0.30196 0.72157 0.33333 scn 5.5074 0 TD [(Plant)-301.3(Science)]TJ 58.8564 0 TD (frontiersin.org)Tj 0.15686 0.15294 0.16078 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[(T)15.6(h)0(e)21(s)0(e)-435.7(s)0(i)19.3(g)0(n)22.6(a)0(l)25.6(s)-448.6(re)24.1(gu)20.5(la)19.3(te)-434.7(no)26.8(du)19.5(la)19.3(ti)20.3(o)15(n)-448(a)15(n)0(d)-438.2(p)0(h)24.2(o)0(s)19.9(p)0(h)24.2(a)0(t)20.9(e)-447(up)19.5(ta)27.2(ke)16.8(,)-443.4(a)0(n)20.5(d)-443.8(ac)21.5(t)]TJ 0 -1.3858 TD [(sy)22(st)23.5(e)12.8(m)0(i)20(c)12.9(al)19.3(ly)-299.9(th)20.4(ro)26.2(ug)20.5(h)-313.1(t)0(h)26.7(e)-308.4(ph)17.9(lo)25.6(em)-302.7(\()]TJ 0.30196 0.72157 0.33333 scn 13.5811 0 TD -0.0132 Tc [(Ya)-13.2(n)7.3(g)-323.7(et)-322.2(a)-13.2(l)6.1(.)-13.2(,)-314.2(2024)]TJ 0.15686 0.15294 0.16078 scn 6.6268 0 TD 0 Tc [(\).)-295.7(Th)17.4(e)12.9(i)0(r)-295.7(m)0(o)20.7(b)0(i)16.7(l)16.9(it)20.3(y)]TJ -20.2079 -1.3921 TD [(st)23.5(ro)19.9(ng)22.6(ly)-344(su)21(gg)21.5(es)17.8(ts)-341.9(th)20.4(at)-344.5(EV)18.1(s)-360.4(c)12.9(ou)24.7(ld)-351.3(a)14.9(c)0(t)-346.6(a)0(s)-351.8(c)12.9(ar)19.9(ri)19.3(e)12.9(r)0(s)-349.2(o)15(f)-357.8(t)0(h)20.4(e)12.9(se)-347.6(re)17.8(gu)20.5(la)25.6(to)20.9(ry)]TJ 0 -1.3858 TD [(RN)17.6(As)-492.3(acr)13.9(os)13.6(s)-499(c)0(e)13.1(l)0(l)-495.3(b)0(o)17.3(und)12.5(ar)13.6(ie)14.6(s)-499(and)-486.2(po)12.1(ssi)17.9(bl)13(y)-499.5(int)19.5(o)-501.6(t)0(h)14.1(e)-497.4(rhi)14.9(zo)15.2(sph)16.6(er)11.5(e,)]TJ ET 0.63529 0.63922 0.63529 scn 49.606 402.293 m 545.953 402.293 l 545.726 402.066 l 49.833 402.066 l f* 545.953 402.293 m 545.953 84.132 l 545.726 84.416 l 545.726 402.066 l f* 545.953 84.132 m 49.606 84.132 l 49.833 84.416 l 545.726 84.416 l f* 49.606 84.132 m 49.606 402.293 l 49.833 402.066 l 49.833 84.416 l f* q 85.493 389.537 m 510.066 389.537 l 510.066 163.446 l 85.493 163.446 l W n q 424.573151 0 0 226.091309 85.492874 163.445648 cm /Im2 Do Q Q BT /F6 1 Tf 5.5001 0 0 5.5001 60.8315 150.9165 Tm 0.43529 0.43529 0.43529 scn [(FIGURE)-305.5(1)]TJ 6.9999 0 0 6.9999 60.8315 141.9024 Tm (\(A\))Tj /F3 1 Tf 1.5469 0 TD [(Conceptual)-300.2(model)-305.2(of)-297(rhizosphere)-304.2(interactions)-306.7(where)-302.4(extracellular)-304.1(vesicles)-300.2(\(EVs\))-303.4(deliver)-300.3(secondary)-301.9(metabolites,)-304.5(proteins,)-303.9(and)-300.2(small)-301.3(RNAs)-303.5(\(sRNAs\))]TJ -1.5469 -1.2878 TD [(to)-308.4(diverse)-296.7(microorganisms.)-307.1(Arrows)-302.5(indicate)-303.4(direct)-299.5(ef)62.2(fects)-303.4(on)-301.4(plant)]TJ /F4 1 Tf 29.821 0 TD ()Tj /F3 1 Tf 0.6965 0 TD [(microbe)-303.1(interactions)-306.7(and)-300.2(indirect)-297.4(ef)62.2(fects)-303.4(via)-304.7(shifts)-301.3(in)-297.6(microbial)-306.3(community)]TJ -30.5175 -1.2797 TD [(composition)-310.5(or)-295.5(nutrient)-305.3(status.)]TJ /F6 1 Tf 14.2869 0 TD (\(B\))Tj /F3 1 Tf 1.5388 0 TD [(Proposed)-302.8(ef)62.2(fects)-303.4(of)-305.1(EV-mediated)-299.2(interactions)-306.7(on)-301.4(nitrogen)-302.6(use)-298.9(ef)]TJ /F5 1 Tf 29.3189 0 TD ()Tj /F3 1 Tf 0.5426 0 TD [(ciency)-296.6(\(NUE\),)-303.4(linking)-304.7(biological)-303.3(nitrogen)]TJ /F5 1 Tf 18.4012 0 TD ()Tj /F3 1 Tf 0.5426 0 TD (xation,)Tj -64.6311 -1.2878 TD [(mineralization,)-306.9(nitri)]TJ /F5 1 Tf 8.6499 0 TD ()Tj /F3 1 Tf 0.5426 0 TD [(cation,)-296(and)-308.3(denitri)]TJ /F5 1 Tf 8.1882 0 TD ()Tj /F3 1 Tf 0.5426 0 TD [(cation)-300.1(processes.)-308(NH)]TJ 5.2498 0 0 4.6661 253.9275 118.0347 Tm (4+)Tj 6.9999 0 0 6.9999 262.7716 114.9165 Tm (from)Tj /F6 1 Tf 2.4864 0 TD (\(B\))Tj /F3 1 Tf 1.5388 0 TD [(is)-297.8(connected)-309.7(to)-300.3(NH)]TJ 5.2498 0 0 4.6661 353.2534 118.0347 Tm (4+)Tj 6.9999 0 0 6.9999 362.0975 114.9165 Tm [(shown)-296.5(in)]TJ /F6 1 Tf 4.4221 0 TD (\(A\))Tj /F3 1 Tf 1.2392 0 TD [(.)-295.5(Primary)-306.9(metabolites)-300.6(\(e.g.,)-307.8(sugars,)-302.1(amino)]TJ -48.7001 -1.2878 TD [(acids,)-303.9(organic)-306.5(acids\))-302.4(are)-301(not)-300.4(shown,)-300.5(as)-300.3(their)-305.9(exudation)-302.6(is)-297.8(largely)-309(independent)-301.8(from)-300.5(EV)-298.1(release)-302.9(and)-300.2(their)-305.9(general)-298.9(roles)-298.6(in)-305.7(the)-297.6(rhizosphere)-304.2(are)-301(well)]TJ T* [(established,)-311.6(which)-302.1(is)-297.8(outside)-305.6(the)-297.6(speci)]TJ /F5 1 Tf 17.324 0 TD ()Tj /F3 1 Tf 0.5426 0 TD [(c)-301.3(focus)-307.3(of)-297(this)-305.1(paper.)]TJ /F9 1 Tf 9.8647 0 TD [(Created)-302.9(in)-303.8(BioRender.)-298.6(Schulz,)-305.3(S.)-299.1(\(2025\))]TJ /F3 1 Tf 0.30196 0.72157 0.33333 scn 18.0692 0 TD (https://BioRender.com/v6i8jao)Tj 0.43529 0.43529 0.43529 scn 13.9953 0 TD (.)Tj /F1 1 Tf 0.49804 0.49804 0.49412 scn -61.3995 99.5383 TD [(Paponov)-303.2(et)-300.8(al.)]TJ 58.5567 0 TD (10.3389/fpls.2025.1681793)Tj -58.5567 -108.3259 TD [(Frontiers)-304.6(in)]TJ 0.30196 0.72157 0.33333 scn 5.5074 0 TD [(Plant)-301.3(Science)]TJ 58.8564 0 TD (frontiersin.org)Tj 0.15686 0.15294 0.16078 scn -29.5213 0 TD (04)Tj ET endstream endobj 81 0 obj<>stream Adobed      ""F   s!1AQa"q2B#R3b$r%C4Scs5D'6Tdt& EFVU(eufv7GWgw8HXhx)9IYiy*:JZjzm!1AQa"q2#BRbr3$4CS%cs5DT &6E'dtU7()󄔤euFVfvGWgw8HXhx9IYiy*:JZjz ?N*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*Ӛ~UUثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWb]v*UثWbw6 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[(reg)-9.2(u)-9.1(lato)-10.2(r)-7.7(y)-505.8(RN)-7.6(As)-511.2(into)-513.5(EVs)-518.7(coul)-14.8(d)-506.8(syn)-10.2(chr)-11.8(oni)-9.2(ze)-509.7(in)-11.6(ter)-7.8(n)-7(a)0(l)-509.8(n)0(u)-9.9(trie)-12.3(nt)]TJ 0 -1.3858 TD [(signaling)-311.3(with)-308.7(microbial)-308(modulation)-315.4(in)-307.7(the)-306.9(rhizosphere,)-315.7(providing)]TJ 0 -1.3921 TD [(plants)-250.5(with)-252(a)-249.6(powerful)-247.8(tool)-249.6(to)-250(dynamically)-250.7(shape)-251.7(microbial)-251.4(activity.)]TJ 1.5559 -1.3858 TD 0.0338 Tc [(Future)-532.9(r)1(esearch)-537.6(s)0.9(hould)-529.7(f)-2.8(o)4.7(c)-3.7(us)-528.2(on)-533.9(quantifying)-535(t)1.9(he)-532.9(a)4.7(c)-3.7(tual)]TJ -1.5559 -1.3921 TD 0 Tc [(contribution)-411.3(of)-405.8(EVs)-411.6(to)-407.4(plant)]TJ /F11 1 Tf 11.9118 0 TD ()Tj /F10 1 Tf 0.5102 0 TD [(microbe)-410.7(communication,)-413(clarifying)]TJ -12.422 -1.3858 TD [(how)-244.7(EV-mediated)-247.3(delivery)-246.9(interacts)-245.2(with)-239.4(other)-242.9(transport)-249.3(pathways,)]TJ 0 -1.3921 TD [(and)-416.9(identifying)-417.4(plant)-413(sRNAs)-419.4(that)-412.2(directly)-421.8(target)-410.7(microbial)-421.4(genes)]TJ 0 -1.3858 TD 0.0187 Tc [(involved)-519.6(in)-517.5(nitrogen)-517.5(cycling.)-512.9(Importan)5.4(tly,)-519.2(several)-512.4(m)-0.8(iRNAs)]TJ /F11 1 Tf 25.8709 0 TD 0 Tc ()Tj /F10 1 Tf -25.8709 -1.3921 TD 0.0365 Tc [(in)4.3(cluding)-532.3(m)4.4(i)0.4(R16)5.3(5)-1(,)-532.9(m)4.4(iR16)5.3(7,)-532.9(m)4.4(i)0.4(R31)5.3(9)-1(,)-532.9(m)-1.9(i)6.7(R)1.7(396)5.3(,)-532.9(m)-1.9(i)6.7(R)-4.6(3)5.3(9)-1(9)5.3(,)-532.9(and)]TJ 0 -1.3858 TD 0.0178 Tc [(miR82)5.5(7)]TJ /F11 1 Tf 3.2252 0 TD 0 Tc ()Tj /F10 1 Tf 0.9323 0 TD 0.0171 Tc [(have)-518.1(also)-516(been)-519.1(show)-5.1(n)-512.8(t)-2.2(o)-516(b)0.6(e)-518.1(d)1.7(if)62.4(ferentially)-520.2(regulated)]TJ -4.1575 -1.3921 TD 0 Tc [(un)-9.9(de)-8.9(r)-505.3(d)0(r)-10.5(oug)-8.6(ht)-508.7(an)-11(d)-506.8(c)0(o)-10(mbi)-9.1(n)-7(e)0(d)-506.5(n)-7(itr)-12.6(oge)-11.8(n)-504.7(d)0(e)]TJ /F14 1 Tf 17.9843 0 TD ()Tj /F10 1 Tf 0.5354 0 TD [(ci)-10.6(ency)-518.6(and)-511.4(d)-9.1(rou)-8.1(ght)]TJ -18.5197 -1.3858 TD [(st)-8(r)-7.7(ess)-8.7(,)-512.7(w)0(i)-7.9(t)0(h)-508.7(d)-9.1(ist)-12.6(inc)-11.3(t)-504.3(e)0(a)-10(r)-7.7(ly)-507.7(\()-9.8(7-d)-7.6(ay)-12.1(\))-507.4(a)0(n)-11(d)-506.8(lat)-12.6(e)-510(\()0(1)-9.5(4-d)-7.6(ay)-12.1(\))-507.4(e)0(x)-10.5(p)0(r)-10.5(ess)-8.7(io)-8.5(n)]TJ T* [(responses)-484.8(\()]TJ 0.30196 0.72157 0.33333 scn 4.6803 0 TD (Yas)Tj /F13 1 Tf 1.4236 0 TD ()Tj /F10 1 Tf T* [(ar)-484(et)-478.8(al.,)-483.3(2024)]TJ 0.15686 0.15294 0.16078 scn 6.0031 0 TD [(\).)-484.7(This)-479.8(temporal)-480.9(plasticity)-487.7(suggests)]TJ -12.1071 -1.3921 TD 0.0416 Tc [(that)-538.3(p)-5.3(l)1.8(ants)-545.6(dyna)-6.4(mically)]TJ /F14 1 Tf 11.3386 0 TD 0 Tc ()Tj /F10 1 Tf 0.5732 0 TD 0.0438 Tc [(ne-t)5.6(un)5.3(e)-541.8(t)-0.7(he)6.3(ir)-543.4(re)6.3(gulat)5.6(o)2.1(ry)-537.6(miRNA)]TJ -11.9118 -1.3858 TD 0 Tc [(expression)-303.5(based)-307.9(on)-300.7(stress)-301.6(type)-300.8(and)-303.6(duration.)]TJ 1.5559 -1.3921 TD [(Emerging)-380.9(evidence)-380.6(from)-376.3(tomato)-372.2(fruit)-379.5(tissue)-381.9(provides)-377.5(indirect)]TJ -1.5559 -1.3858 TD [(but)-410.3(compelling)-421.1(support)-412.3(for)-413.5(this)-413.9(hypothesis.)-417.8(Under)-415.8(salt,)-413.2(drought,)]TJ 0 -1.3921 TD [(and)-240.6(nitrogen)-246.3(de)]TJ /F14 1 Tf 6.3055 0 TD ()Tj /F10 1 Tf 0.5291 0 TD [(ciency)-245.7(stress,)-241(tomato)-239.9(plants)-244.2(not)-244.4(only)-241.5(increased)-240.8(EV)]TJ -6.8346 -1.3858 TD [(secretion)-358(but)-353.6(also)-357.6(selectively)-353(modulated)-359.2(the)-357.3(miRNA)-352.5(composition,)]TJ 0 -1.3921 TD [(enriching)-332.7(miR162)-322.6(and)-328.8(miR1919)-328.7(while)-322.7(depleting)-335.9(miR9476)-322.4(\()]TJ 0.30196 0.72157 0.33333 scn 24.0693 0 TD (Huang)Tj -24.0693 -1.3858 TD [(et)-522.9(al)-12.2(.,)-521.5(20)-12.1(25)]TJ 0.15686 0.15294 0.16078 scn 4.8378 0 TD [(\).)-522.5(Th)-7.8(e)-12.3(s)-7.7(e)-510(s)-7.7(hi)-15.3(ft)-11.7(s)-511.6(w)-9.6(er)-13.7(e)-510(s)-7.7(t)-6.7(r)-7.7(es)-13.7(s)-7.7(-)0(s)-12.7(p)-9.1(ec)-12.1(i)]TJ /F14 1 Tf 13.6126 0 TD ()Tj /F10 1 Tf 0.5417 0 TD 0.0074 Tc [(c)-502.6(a)-2.8(nd)-505.7(f)7.4(u)-6.8(nc)7.4(t)-5.3(i)7.4(o)-7.4(n)0.4(a)7.4(l)-4.8(ly)]TJ -18.9921 -1.3921 TD 0.0528 Tc [(consequenti)4.1(a)-1.5(l:)-548.1(ch)4.3(anges)-553.3(i)-2.2(n)-552.7(E)3.8(V)-554.1(c)2.7(argo)-549.6(directl)6.7(y)-553.8(altered)-548.5(z)-3.6(i)4.1(n)1.7(c)]TJ 0 -1.3858 TD 0 Tc [(transporter)-464.5(gene)-461.9(expression)-467.3(and)-467.3(zinc)-458.3(accumulation)-466.1(in)-465.1(intestinal)]TJ 0 -1.3921 TD [(cells.)-252.9(Although)-258.8(derived)-252.3(from)-256.6(fruit)-253.5(tissues)-257.2(rather)-250.5(than)-255.1(root)-248.9(exudates,)]TJ 0 -1.3858 TD 0.015 Tc [(the)-3.6(s)1(e)]TJ /F14 1 Tf 2.6268 0 TD 0 Tc ()Tj /F10 1 Tf 0.548 0 TD 0.0159 Tc [(nding)-521.4(s)1.9(trongly)-521.4(s)1.9(uggest)-513.6(that)-513.6(plants)-520.9(can)-520.3(r)1.9(eprogra)5.7(m)-520.1(EV)]TJ -3.1748 -1.3921 TD 0 Tc [(composition)-429.7(in)-421(response)-426.8(to)-426.3(abiotic)-424.2(stress)]TJ /F11 1 Tf 17.0457 0 TD ()Tj /F10 1 Tf 0.9134 0 TD [(raising)-427.8(the)-426.6(possibility)]TJ -17.9591 -1.3858 TD [(that)-324(similar)-327(mechanisms)-325.7(operate)-326.9(in)-320.3(root-derived)-329.9(EVs)-323.4(to)-325.6(modulate)]TJ 0 -1.3921 TD [(microbial)-301.7(activity)-307(in)-301.4(the)-300.6(rhizosphere.)]TJ 1.5559 -1.3858 TD [(If)-497.3(these)-503.3(regulatory)-497.9(miRNAs)-505.1(are)-496.4(indeed)-501.8(packaged)-502.2(into)-500.9(EVs,)]TJ -1.5559 -1.3921 TD [(plants)-401.7(may)-403.2(be)-400.5(capable)-406.7(of)-399.5(time-sensitive)-408.8(modulation)-403.6(of)-399.5(microbial)]TJ 0 -1.3858 TD [(activity)-319.6(in)-314(the)-319.5(rhizosphere,)-322(adapting)-317.1(their)-319.1(in)]TJ /F14 1 Tf 17.9024 0 TD ()Tj /F10 1 Tf 0.5291 0 TD [(uence)-324(in)-314(response)-319.7(to)]TJ -18.4315 -1.3921 TD [(both)-359.1(immediate)-357.1(and)-360.3(prolonged)-359.3(environmental)-360.4(challenges.)-364.4(To)-360(test)]TJ 0 -1.3858 TD [(these)-396.2(hypotheses,)-406.7(several)-400.5(methodological)-399.1(and)-398.1(analytical)-400.9(advances)]TJ 0 -1.3921 TD [(are)-357.8(critical)]TJ /F11 1 Tf 4.2835 0 TD ()Tj /F10 1 Tf 0.9134 0 TD [(particularly)-360.5(in)-351.8(isolating)-357.1(EVs)-354.9(from)-357.4(soil)-352(environments,)]TJ -5.1969 -1.3858 TD [(characterizing)-227.6(their)-218.3(cargo,)-221.2(and)-221.7(linking)-218.3(these)-219.8(molecules)-226(to)-218.5(functional)]TJ 0 -1.3921 TD [(outcomes)-346.3(in)-339.2(microbial)-345.8(communities.)-342.7(Complementary)-350.6(approaches,)]TJ 0 -1.3858 TD [(inc)-11.3(ludi)-12.2(ng)-506.5(m)-6.9(etag)-12.2(enom)-11.3(ics)-509.6(an)-11(d)-506.8(trans)-14(crip)-14.8(tomi)-9.4(cs,)-513.8(can)-508.4(he)-10.5(lp)-508.7(iden)-14.2(tify)]TJ 0 -1.3921 TD [(crop)-404.7(genotypes)-402.9(naturally)-403.2(predisposed)-401.3(to)-401.1(associate)-403.2(with)-403.2(microbial)]TJ 0 -1.3858 TD [(consortia)-303.4(that)-298.9(enhance)-305.1(nitrogen-use)-305.7(ef)]TJ /F14 1 Tf 15.452 0 TD ()Tj /F10 1 Tf 0.5291 0 TD (ciency.)Tj -14.4252 -1.3921 TD [(In)-316.5(parallel,)-322.5(EV)-315.8(research)-320.3(in)-314(plants,)-322.3(particularly)-322.7(their)-312.8(role)-322(in)-314(the)]TJ -1.5559 -1.3858 TD [(rhizosphere,)-416.5(faces)-406.7(signi)]TJ /F14 1 Tf 9.5307 0 TD ()Tj /F10 1 Tf 0.5354 0 TD [(cant)-407.9(bioinformatics)-410.3(and)-410.7(data)-407.9(integration)]TJ -10.0661 -1.3921 TD [(challen)-12.5(ges.)-503(Isol)-8.1(ating)-502.8(and)-505.1(cha)-8.1(racteri)-11.1(zing)-504.5(EVs)-506.1(from)-508.6(complex)-510.9(soil)]TJ 0 -1.3858 TD [(environments)-481.8(is)-484.7(a)-476.4(major)-477.9(hurdle,)-488.1(as)-477.7(plant-derived)-487.6(EVs)-474.6(must)-483.9(be)]TJ 0 -1.3921 TD [(distinguished)-372.1(from)-376.3(microbial)-371(and)-366.6(abiotic)-373.8(particles.)-371.1(Once)-369.9(isolated,)]TJ 28.3654 73.6126 TD [(EVs)-512.4(con)-10.7(tai)-8.9(n)-504.7(a)-507.9(hi)-9(ghl)-8.2(y)-505.8(div)-9.2(ers)-8.7(e)-510(car)-11.3(go)]TJ /F11 1 Tf 15.1307 0 TD ()Tj /F10 1 Tf 0.9197 0 TD [(inc)-11.3(ludi)-12.2(n)-7(g)-505.8(pro)-8.1(tei)-11(ns)-8.4(,)-506.4(lip)-9.4(ids)-8.7(,)]TJ -16.0504 -1.3858 TD 0.0376 Tc [(metabolites,)-544.4(and)-538.5(R)-3.5(NA)-6.1(s)]TJ /F11 1 Tf 10.4819 0 TD 0 Tc ()Tj /F10 1 Tf 0.9575 0 TD 0.0372 Tc [(whic)-6.6(h)-534.2(r)-2(eq)-7.1(ui)-5.2(res)-537.4(t)-1(h)-5(e)-535.8(a)-4.5(p)2.9(plica)-4.5(t)-1(io)-4.5(n)-536.8(o)-4.5(f)]TJ -11.4394 -1.3921 TD 0 Tc [(m)-6.9(u)-9.1(l)-8.3(t)-6.7(i)-10.9(p)-9.1(l)-8.3(e)-516.3(o)-10.2(m)-6.9(i)-10.9(c)0(s)-523.9(a)-10.3(p)-9.1(p)-9.1(r)-7.7(o)-10.2(a)0(c)-16.3(h)-10.7(es)-523.9(s)-7.7(u)-9.1(c)-12.3(h)-514.7(a)-10.2(s)-517.9(p)-9.1(r)-7.7(o)-10.2(t)-6.7(eo)-16.3(m)-6.9(i)-10.9(c)-12.3(s)-7.7(,)-512.7(m)-13.2(et)-19(ab)-14.2(o)-10.2(l)-8.3(om)-17.1(ic)-16.9(s)-7.7(,)]TJ 0 -1.3858 TD 0.0248 Tc [(an)5.2(d)-519.8(s)-1.8(m)5.3(a)2(ll)-518.9(RNA)-522.8(s)4.5(equ)3.1(e)-0.1(n)5.2(c)-0.1(in)5.2(g.)-519.4(In)5.2(te)6.2(gr)4.5(atin)5.2(g)-525.1(t)5.5(hes)4.6(e)-523(het)5.5(e)-0.1(r)4.6(o)2(gen)5.2(e)-0.1(ou)3.1(s)]TJ 9 0 0 9 304.8944 702.7085 Tm 0.0361 Tc [(da)7(tas)3.2(e)4.9(ts)-532.2(t)4.2(o)-528.5(in)3.9(fer)-532.2(t)4.2(he)-530.6(fun)3.9(c)4.9(ti)6.3(ona)7(l)-532.8(r)3.2(o)0.7(le)-530.6(of)-535.9(E)6(V)2.4(s)-532.2(i)0(s)-532.2(p)1.8(ar)3.2(ti)6.3(cular)3.3(l)2.6(y)]TJ T* 0 Tc [(challenging,)-332.6(especially)-338.9(when)-329.5(attempting)-336.6(to)-331.9(correlate)-333.4(speci)]TJ /F14 1 Tf 23.326 0 TD ()Tj /F10 1 Tf 0.5354 0 TD [(c)-327.3(cargo)]TJ -23.8614 -1.3921 TD [(molecules)-301.6(\(e.g.,)-308.5(miRNAs)-303.5(or)-307.7(secondary)-305.2(metabolites\))-307.4(with)-302.4(microbial)]TJ 0 -1.3858 TD [(community)-298.6(responses.)]TJ 1.5496 -1.3921 TD [(T)-9.6(h)0(e)-508.1(s)0(p)-10.5(ati)-8.9(al)-509.8(and)-517.7(temp)-9.8(or)-11.6(al)-509.8(dyn)-11.7(ami)-9.1(cs)-511.3(of)-512.9(EV)-511.1(sec)-13.4(ret)-7.8(io)-8.5(n)-504.7(u)-9.1(nde)-9.6(r)]TJ -1.5496 -1.3858 TD [(changing)-452.5(environmental)-454.9(conditions)-451.1(further)-449.8(complicate)-452.1(this)-451.6(task.)]TJ 0 -1.3921 TD [(In)-222.1(this)-224.9(context,)-225.9(multivariate)-233.9(techniques)-227.9(that)-223.3(explore)-228.2(the)-225(latent)-227.6(space)]TJ /F11 1 Tf 0 -1.3858 TD ()Tj /F10 1 Tf 0.9512 0 TD 0.0401 Tc [(wh)4.2(en)-540.2(cou)5.8(p)-0.5(led)-536(w)-1(i)4(t)1.9(h)-537.6(m)1.7(ech)4.2(a)-1.6(ni)4(stic)-532.9(know)5.3(ledge)-539.2(o)4.7(f)-538.2(m)1.7(ol)6.6(ecul)6.6(ar)]TJ -0.9512 -1.3921 TD 0 Tc 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[(ma)27(nu)15.3(f)13.8(a)0(c)21.5(t)0(u)22(r)0(e)17.8(r)0(,)-274.7(i)0(s)-283.1(n)0(o)20.5(t)-283.8(gu)20.5(ar)19.9(an)20.5(te)18.8(e)12.9(d)-292.6(o)15(r)-291.1(e)12.9(n)0(d)21.6(o)0(r)19.9(s)0(e)17.8(d)-286.3(by)-276.6(th)20.4(e)-289.5(p)0(u)19.5(b)0(l)19.3(i)14.3(sh)19.4(e)12.8(r)0(.)]TJ /F1 1 Tf 13.5 0 0 13.5 49.6063 581.2156 Tm 0.30196 0.72157 0.33333 scn (References)Tj /F10 1 Tf 6.9999 0 0 6.9999 56.126 556.2141 Tm 0.13725 0.12157 0.12549 scn [(Ambastha,)-215.2(V.,)-211.5(Nevo,)-220.3(Y.,)-210.6(Matityhu,)-217.7(I.,)-219.1(Honys,)-212.5(D.,)-220(and)-210.9(Leshem,)-219(Y.)-218(\(2023\).)-209.2(Discovery)-222.4(of)]TJ -0.9314 -1.142 TD [(microRNAs)-419.5(in)]TJ /F15 1 Tf 6.3983 0 TD [(Pyrus)-415.3(stigma)]TJ /F10 1 Tf 5.6451 0 TD [(exudates)-420.4(opens)-420.9(new)-415.1(research)-425.8(avenues)-419.8(in)-422(Horticulture.)]TJ /F15 1 Tf -12.0434 -1.142 TD [(PNAS)-301.7(Nexus)]TJ /F10 1 Tf 5.394 0 TD [(2.)-299.2(doi:)-300(10.1093/pnasnexus/pgad332)]TJ -4.4626 -1.5712 TD [(Ambrosone,)-208.7(A.,)-205.3(Barbulova,)-203.8(A.,)-205.3(Cappetta,)-209.3(E.,)-208.8(Cillo,)-206.7(F.,)-202.2(De)-207.6(Palma,)-207.9(M.,)-201.7(Ruocco,)-209.6(M.,)-209.8(et)-205.3(al.)]TJ -0.9314 -1.142 TD [(\(2023\).)-427.8(Plant)-435(extracellular)-431.3(vesicles:)-434.1(Current)-433.1(landscape)-432.1(and)-437.7(future)-429.9(directions.)]TJ /F15 1 Tf 31.9916 0 TD (Plants)Tj -31.9916 -1.142 TD (\(Basel\))Tj /F10 1 Tf 2.9886 0 TD [(12.)-298.1(doi:)-300(10.3390/plants12244141)]TJ -2.0572 -1.5712 TD (Ayala-Garc)Tj /F12 1 Tf 4.5517 0 TD ()Tj /F13 1 Tf 0.2916 0.0405 TD ()Tj /F10 1 Tf -0.0324 -0.0405 TD [(a,)-470.8(P.,)-476.1(Herrero-Go)]TJ /F13 1 Tf 7.2649 0.0405 TD ()Tj /F10 1 Tf 0.081 -0.0405 TD [(mez,)-478.7(I.,)-470.2(Jime)]TJ /F13 1 Tf 5.3535 0.0405 TD ()Tj /F10 1 Tf 0.0405 -0.0405 TD [(nez-Guerrero,)-477.5(I.,)-478.3(Otto,)-466(V.,)-478.8(Moreno-de)]TJ -18.4822 -1.142 TD [(Castro,)-390.6(N.,)-396.3(Msken,)-398(M.,)-387.9(et)-391.6(al.)-399(\(2025\).)-387.3(Extracellular)-398.7(vesicle-driven)-395.2(crosstalk)-401.7(between)]TJ 0 -1.1501 TD [(legume)-448.7(plants)-445.1(and)-445.8(rhizobia:)-442.2(The)-447.6(peribacteroid)-443.1(space)-446.9(of)-448.2(symbiosomes)-441(as)-446.3(a)-445.8(protein)]TJ 0 -1.142 TD (traf)Tj /F14 1 Tf 1.3364 0 TD ()Tj /F10 1 Tf 0.5264 0 TD [(cking)-306.3(interface.)]TJ /F15 1 Tf 6.4388 0 TD [(J.)-293.2(Proteome)-308.5(Res.)]TJ /F10 1 Tf 6.7223 0 TD [(24,)-298.1(94)]TJ /F11 1 Tf 2.4297 0 TD ()Tj /F10 1 Tf 0.5102 0 TD [(110.)-305(doi:)-300(10.1021/acs.jproteome.4c00444)]TJ -17.0325 -1.5712 TD [(Badri,)-323.2(D.)-332.7(V.,)-324.9(and)-332.4(Vivanco,)-328.3(J.)-326.6(M.)-330.6(\(2009\).)-330.6(Regulation)-326.7(and)-332.4(function)-330.4(of)-326.7(root)-326.4(exudates.)]TJ /F15 1 Tf -0.9314 -1.142 TD [(Plant)-300.6(Cell)-300(Environ.)]TJ /F10 1 Tf 7.8967 0 TD [(32,)-306.2(666)]TJ /F11 1 Tf 2.9157 0 TD ()Tj /F10 1 Tf 0.5102 0 TD [(681.)-305(doi:)-300(10.1111/j.1365-3040.2009.01926.x)]TJ -10.3912 -1.5712 TD [(Baldrich,)-292.5(P.,)-297.9(Rutter,)-293.1(B.)-292.4(D.,)-292.9(Karimi,)-295.6(H.)-293.6(Z.,)-291.2(Podicheti,)-299.6(R.,)-294.5(Meyers,)-296.9(B.)-292.4(C.,)-293(and)-291.9(Innes,)-300.3(R.)]TJ -0.9314 -1.142 TD [(W.)-458.2(\(2019\).)-468.3(Plant)-459.3(extracellular)-463.7(vesicles)-465.8(contain)-461.4(diverse)-466(small)-462.6(RNA)-454.6(species)-467.5(and)-462(are)]TJ T* [(enriched)-367(in)-357.2(10-)-356.5(to)-364.3(17-nucleotide)]TJ /F11 1 Tf 13.6389 0 TD ()Tj /F10 1 Tf 0.3969 0 TD (tiny)Tj /F11 1 Tf 1.5631 0 TD ()Tj /F10 1 Tf 0.7532 0 TD (RNAs.)Tj /F15 1 Tf 2.9805 0 TD [(Plant)-365.4(Cell)]TJ /F10 1 Tf 4.333 0 TD [(31,)-362.9(315)]TJ /F11 1 Tf 2.9724 0 TD ()Tj /F10 1 Tf 0.5102 0 TD [(324.)-361.7(doi:)-364.8(10.1105/)]TJ -27.1483 -1.142 TD (tpc.18.00872)Tj 0.9314 -1.5712 TD [(Cai,)-340.4(Q.,)-347.7(Halilovic,)-346(L.,)-345.1(Shi,)-345.3(T.,)-343.1(Chen,)-351.3(A.,)-343(He,)-347.3(B.,)-341.7(Wu,)-343.1(H.,)-351(et)-343(al.)-342.3(\(2023\).)-346.8(Extracellular)]TJ -0.9314 -1.142 TD [(vesicles:)-328.8(cross-organismal)-326.2(RNA)-325(traf)]TJ /F14 1 Tf 14.3031 0 TD ()Tj /F10 1 Tf 0.5264 0 TD [(cking)-322.5(in)-324.8(plants,)-324.3(microbes,)-329.3(and)-324.3(mammalian)-328.9(cells.)]TJ /F15 1 Tf -14.8295 -1.142 TD [(Extracellular)-307.7(vesicles)-300.7(circulating)-308.8(Nucleic)-301(Acids)]TJ /F10 1 Tf 18.5227 0 TD [(4,)-299.2(262.)-305(doi:)-300(10.20517/evcna.2023.10)]TJ -17.5913 -1.5712 TD [(Cai,)-218.9(Q.,)-226.2(Qiao,)-227.2(L.,)-223.6(Wang,)-220.9(M.,)-226(He,)-225.9(B.,)-220.2(Lin,)-223.7(F.-M.,)-227.4(Palmquist,)-227.7(J.,)-222(et)-229.6(al.)-220.8(\(2018\).)-225.3(Plants)-224.9(send)]TJ -0.9314 -1.1501 TD [(small)-430.2(RNAs)-430.8(in)-438.2(extracellular)-431.3(vesicles)-433.4(to)-429.1(fungal)-435(pathogen)-433.7(to)-429.1(silence)-439(virulence)-433(genes.)]TJ /F15 1 Tf 0 -1.142 TD (Science)Tj /F10 1 Tf 3.102 0 TD [(360,)-305(1126)]TJ /F11 1 Tf 3.8714 0 TD ()Tj /F10 1 Tf 0.5102 0 TD [(1129.)-303.9(doi:)-300(10.1126/science.aar4142)]TJ -6.5522 -1.5712 TD [(Canarini,)-408.6(A.,)-407.8(Kaiser,)-409.4(C.,)-406.4(Merchant,)-409.4(A.,)-399.7(Richter,)-410.3(A.,)-407.8(and)-405.3(Wanek,)-413.7(W.)-401.5(\(2019\).)-411.6(Root)]TJ -0.9314 -1.142 TD [(exudation)-352.9(of)-359.1(primary)-348.7(metabolites:)-358(Mechanisms)-360.1(and)-348.6(their)-354.3(roles)-357.6(in)-349.1(plant)-355.5(responses)-354.6(to)]TJ T* [(environmental)-309.4(stimuli.)]TJ /F15 1 Tf 9.4355 0 TD [(Front.)-299.3(Plant)-300.6(Sci.)]TJ /F10 1 Tf 6.7628 0 TD [(10.)-298.1(doi:)-308.1(10.3389/fpls.2019.00157)]TJ -15.2669 -1.5712 TD [(Glick,)-356.1(B.)-365.3(R.)-358.6(\(2014\).)-363(Bacteria)-364.9(with)-360.1(ACC)-357.9(deaminase)-365(can)-363.5(promote)-360.3(plant)-363.6(growth)-359.1(and)]TJ -0.9314 -1.142 TD 0.0643 Tc [(h)-4(e)-5.6(lp)-573.9(to)-569.6(f)-9.2(e)2.4(e)-5.6(d)-573.9(th)-4(e)-572.6(w)-3.8(orl)-9.7(d)1.2(.)]TJ /F15 1 Tf 12.0677 0 TD 0.0653 Tc [(Mi)-4.9(cr)-8.1(obi)-4.9(o)-1.2(log)-4.6(i)-4.9(c)-5.2(a)3.6(l)-570.4(R)-5.8(es)-6.7(.)]TJ /F10 1 Tf 10.0348 0 TD 0.0717 Tc [(16)8.1(9,)-560.7(30)]TJ /F11 1 Tf 3.5879 0 TD 0 Tc ()Tj /F10 1 Tf 0.575 0 TD 0.0644 Tc [(3)-7.3(9)0.8(.)-576.1(d)1.3(oi)-7.7(:)-568(1)-7.3(0.)-9.2(10)-7.3(16)-7.3(/)]TJ -26.2655 -1.142 TD 0 Tc (j.micres.2013.09.009)Tj 0.9314 -1.5712 TD [(Huang,)-216.6(Z.,)-218.3(Pych,)-220.5(E.,)-225(Whitehead,)-220.1(B.)-219.5(J.,)-222(Nejsum,)-220.4(P.)-216.2(L.,)-223.6(Bellini,)-218.8(I.,)-219.1(Drace,)-222.4(T.,)-221.6(et)-213.4(al.)-220.8(\(2025\).)]TJ -0.9314 -1.142 TD [(Impact)-233.4(of)-237.6(abiotic)-232.7(stress)-231.5(on)-238.7(miRNA)-233(pro)]TJ /F14 1 Tf 15.4046 0 TD ()Tj /F10 1 Tf 0.5345 0 TD [(les)-233.5(in)-227.6(tomato-derived)-237.5(extracellular)-245(vesicles)-230.9(and)]TJ -15.9391 -1.142 TD [(t)-13.9(h)-11.6(ei)-20.5(r)-518.8(b)-18.3(io)-17.5(l)-9.2(o)-18.3(g)-11.8(i)0(c)-20.5(a)0(l)-536(a)0(c)-21.7(t)0(i)-21.1(v)-11.8(it)-21.1(y)-11.7(.)]TJ /F15 1 Tf 10.6261 0 TD [(In)-18.8(t)-13.9(.)-516.1(J)-12(.)-524.2(B)-11.1(i)-13.5(o)-9.8(l)-12.1(.)-524.2(M)0(a)-21(c)-13.8(ro)-26.5(mo)-22.6(l)-12.1(e)-10(c)-13.8(u)-11.6(le)-22(s)]TJ /F10 1 Tf 12.3674 0 TD 0.0224 Tc [(3)7.4(1)22.4(9,)-496.6(1)7.4(4)22.4(52)7.4(6)22.4(0)0.4(.)-496.6(d)7.9(o)12.2(i)22.4(:)-512(1)22.4(0.)22.4(10)22.4(16)22.4(/)]TJ -22.9934 -1.142 TD 0 Tc (j.ijbiomac.2025.145260)Tj 0.9314 -1.5793 TD [(Islam,)-242.6(W.,)-248.3(Tauqeer,)-246.9(A.,)-245.8(Waheed,)-252.1(A.,)-245.8(and)-243.3(Zeng,)-252.4(F.)-242(\(2022\).)-249.6(MicroRNA)-248.8(mediated)-245.2(plant)]TJ -0.9314 -1.142 TD [(responses)-306(to)-299.5(nutrient)-301.5(stress.)]TJ /F15 1 Tf 11.5008 0 TD [(Int.)-305.8(J.)-293.2(Mol.)-302.9(Sci.)]TJ /F10 1 Tf 6.2849 0 TD [(23,)-298.1(2562.)-303.9(doi:)-300(10.3390/ijms23052562)]TJ -16.8543 -1.5712 TD [(Ju,)-244(Y.,)-251.1(Jia,)-245.2(Y.,)-243(Cheng,)-249.6(B.,)-244.5(Wang,)-253.3(D.,)-244.3(Gu,)-246.8(D.,)-244.3(Jing,)-250.1(W.,)-248.3(et)-245.8(al.)-245.1(\(2024\).)-249.6(NRT1.1B)-250.8(mediates)]TJ -0.9314 -1.142 TD [(rice)-220.5(plant)-217.8(growth)-221.4(and)-219(soil)-213.5(microbial)-226.4(diversity)-218.4(under)-222.7(dif)58.6(ferent)-213.2(nitrogen)-222.2(conditions.)]TJ /F15 1 Tf 32.3398 0 TD (AMB)Tj -32.3398 -1.142 TD (Express)Tj /F10 1 Tf 3.2235 0 TD [(14,)-306.2(39.)-298.1(doi:)-300(10.1186/s13568-024-01683-7)]TJ -2.2921 -1.5712 TD [(Kretzschmar,)-282.6(T.,)-278.3(Kohlen,)-279.7(W.,)-272.6(Sasse,)-284.3(J.,)-278.7(Borghi,)-277.1(L.,)-280.3(Schlegel,)-278.4(M.,)-282.7(Bachelier,)-280.2(J.)-278(B.,)-276.9(et)-278.2(al.)]TJ -0.9314 -1.142 TD [(\(2012\).)-314.4(A)-309.2(petunia)-316.6(ABC)-317.5(protein)-312.4(controls)-318.6(strigolactone-dependent)-322.4(symbiotic)-307.3(signalling)]TJ T* [(and)-300(branching.)]TJ /F15 1 Tf 6.374 0 TD (Nature)Tj /F10 1 Tf 3.0777 0 TD [(483,)-305(341)]TJ /F11 1 Tf 3.3935 0 TD ()Tj /F10 1 Tf 0.5102 0 TD [(U135.)-305(doi:)-300(10.1038/nature10873)]TJ -12.4241 -1.5712 TD [(Kuppe,)-242(C.)-243.7(W.,)-248.3(and)-243.3(Postma,)-247.2(J.)-237.5(A.)-245.1(\(2024\).)-249.6(Bene)]TJ /F14 1 Tf 18.0044 0 TD ()Tj /F10 1 Tf 0.5345 0 TD [(ts)-241.1(and)-243.3(limits)-246.6(of)-245.7(biological)-243(nitri)]TJ /F14 1 Tf 11.9786 0 TD ()Tj /F10 1 Tf 0.5345 0 TD (cation)Tj -31.9835 -1.142 TD [(in)-25.1(h)-11.6(i)0(b)-17.5(i)-15.4(to)-16(r)-16.7(s)-518.8(fo)-18.9(r)-526.9(p)0(l)-23.7(a)0(n)-18.2(t)-524.1(n)-9.7(i)-15.4(tr)-22.4(o)-10.2(g)-11.7(en)-533.3(up)-21(ta)-22.3(k)-13.1(e)-515.4(a)-16.6(n)-9.7(d)-524.8(t)0(h)-17.4(e)-523.5(e)-13.3(n)-9.7(v)-11.7(i)0(r)-23.9(o)-10.2(n)-9.7(m)-14.1(en)-23(t.)]TJ /F15 1 Tf 25.9172 0 TD 0.0134 Tc [(Sc)13.4(i)-5.7(.)-510.8(R)13.4(e)-2.9(p.)]TJ /F10 1 Tf 4.1387 0 TD 0.022 Tc [(1)22(4)0(,)-497(1)22(50)22(27)22(.)]TJ -30.0559 -1.142 TD 0 Tc [(doi:)-300(10.1038/s41598-024-65247-2)]TJ 0.9314 -1.5712 TD [(Kuzyakov,)-487(Y.,)-486(and)-486.3(Razavi,)-488.8(B.)-486.8(S.)-480.3(\(2019\).)-492.6(Rhizosphere)-486.5(size)-487.8(and)-486.3(shape:)-486.5(Temporal)]TJ -0.9314 -1.142 TD [(dynamics)-484.6(and)-486.3(spatial)-483.4(stationarity.)]TJ /F15 1 Tf 6.9999 0 0 6.9999 151.4834 162.2551 Tm [(Soil)-484.5(Biol.)-489.8(Biochem.)]TJ /F10 1 Tf 8.2692 0 TD [(135,)-483.2(343)]TJ /F11 1 Tf 3.5717 0 TD ()Tj /F10 1 Tf 0.5102 0 TD [(360.)-483.2(doi:)-486.3(10.1016/)]TJ -26.9053 -1.1501 TD (j.soilbio.2019.05.011)Tj 0.9314 -1.5712 TD [(Lazicki,)-212.7(P.,)-208.8(Geisseler,)-217(D.,)-203.8(and)-210.9(Lloyd,)-212(M.)-209.1(\(2020\).)-217.3(Nitrogen)-210.5(mineralization)-213.7(from)-211.7(organic)]TJ -0.9314 -1.142 TD [(amendments)-346.6(is)-339.8(variable)-344.5(but)-346.5(predictable.)]TJ /F15 1 Tf 16.5708 0 TD [(J.)-341.8(Environ.)-343.7(Qual.)]TJ /F10 1 Tf 7.1515 0 TD [(49,)-338.6(483)]TJ /F11 1 Tf 2.9562 0 TD ()Tj /F10 1 Tf 0.5102 0 TD [(495.)-337.4(doi:)-348.6(10.1002/)]TJ -27.1888 -1.142 TD (jeq2.20030)Tj 0.9314 -1.5712 TD [(Lee)-370.2(D)]TJ /F12 1 Tf 2.4621 0 TD ()Tj /F13 1 Tf 0.2916 0.0405 TD ()Tj /F10 1 Tf -0.0324 -0.0405 TD [(az,)-370.7(A.)-374.7(S.,)-367.6(Rizaludin,)-378.1(M.)-371.1(S.,)-375.7(Zweers,)-369.9(H.,)-375.3(Raaijmakers,)-377.2(J.)-367.1(M.,)-371.8(and)-372.9(Garbeva,)-374(P.)]TJ -3.6527 -1.142 TD [(\(2022\).)-354.9(Exploring)-361.5(the)-354.7(volatiles)-351(released)-364.5(from)-349.4(roots)-359.3(of)-351(wild)-358.3(and)-356.7(domesticated)-355.9(tomato)]TJ T* [(plants)-299.3(under)-303.7(insect)-309(attack.)]TJ /F15 1 Tf 10.8852 0 TD (Molecules)Tj /F10 1 Tf 4.1549 0 TD [(27,)-306.2(1612.)-295.8(doi:)-308.1(10.3390/molecules27051612)]TJ -14.1087 -1.5712 TD [(Liu,)-325.7(S.,)-335.2(Li,)-327.4(H.-Y.,)-336.5(Zhang,)-332.1(T.,)-326.9(Li,)-335.5(Y.-K.,)-325.2(Liao,)-338(X.-D.,)-331.3(and)-332.4(Xing,)-332.1(S.-C.)-329.5(\(2025\).)-330.6(Organic)]TJ -0.9314 -1.142 TD [(fertilizer)-208.5(enhances)-209.5(the)-200.8(secretion)-211.5(of)-205.2(microRNAs)-208.9(from)-203.6(tomato)-208.5(roots)-205.4(to)-202.3(facilitate)-208.9(bene)]TJ /F14 1 Tf 32.5423 0 TD ()Tj /F10 1 Tf 0.5345 0 TD (cial)Tj 3.3935 67.8545 TD 0.0339 Tc [(rhizos)-7.1(phere)-538.2(m)-4.5(icroo)-8.7(r)1(gani)-5.8(sm)-539.1(exp)-4.9(a)1.1(ns)-7.1(ion)-534.7(a)-6.9(nd)-539.5(su)-4.9(pp)-4.9(ress)]TJ /F15 1 Tf 23.9896 0 TD 0.0367 Tc [(Ra)7.4(lst)6.6(o)-5.5(ni)7(a)-535.3(s)-2.9(olana)7.3(c)-1.3(ear)3.8(u)0.8(m)]TJ /F10 1 Tf -23.9896 -1.142 TD 0 Tc (proliferation.)Tj /F15 1 Tf 5.5236 0 TD (Microbiome)Tj /F10 1 Tf 5.0539 0 TD [(13,)-298.1(159.)-305(doi:)-300(10.1186/s40168-025-02137-3)]TJ -9.6542 -1.5712 TD [(Lu,)-205.1(Y.,)-210.6(Hua,)-211.3(Y.,)-202.5(Lv,)-210.4(N.,)-201.9(Zu,)-207.9(W.,)-207.8(Kronzucker,)-216.9(H.)-204.5(J.,)-205.8(Dong,)-210.6(G.,)-208.6(et)-205.3(al.)-204.6(\(2022\).)-209.2(Syringic)-209.1(acid)]TJ -0.9233 -1.142 TD [(from)-276.5(rice)-285.3(roots)-278.3(inhibits)-286.5(soil)-278.3(nitri)]TJ /F14 1 Tf 12.9829 0 TD ()Tj /F10 1 Tf 0.5264 0 TD [(cation)-289.7(and)-275.7(N)]TJ 5.2498 0 0 4.6661 436.3085 528.0377 Tm (2)Tj 6.9999 0 0 6.9999 438.803 529.2283 Tm [(O)-281.5(emission)-281.6(under)-287.5(red)-279.4(and)-283.8(paddy)-282.6(soils)]TJ -19.1302 -1.142 TD [(but)-297.9(not)-301.2(a)-300(calcareous)-302.2(soil.)]TJ /F15 1 Tf 10.3183 0 TD [(Front.)-307.4(Plant)-300.6(Sci.)-296(Volume)]TJ /F10 1 Tf 10.1482 0 TD [(13.)-298.1(doi:)-308.1(10.3389/fpls.2022.1099689)]TJ -19.5432 -1.5712 TD [(Ma,)-411.9(W.,)-410.2(Tang,)-412.9(S.,)-416.2(Dengzeng,)-407.4(Z.,)-412.7(Zhang,)-413.1(D.,)-414.4(Zhang,)-413.1(T.,)-407.9(and)-413.4(Ma,)-411.9(X.)-408.6(\(2022\).)-411.6(Root)]TJ 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Paponov Stefanie Schulz Michael Schloter Ana Conesa Yehoram Leshem Paponov IA, Schulz S, Schloter M, Conesa A and Leshem Y (2025) Extracellular vesicles in the rhizosphere: targets to improve nutrient use efficiency of crops?. Front. Plant Sci. 16:1681793. doi: 10.3389/fpls.2025.1681793 Copyright © 2025 Paponov, Schulz, Schloter, Conesa and Leshem.. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms. extracellular vesicles nutrient use efficiency rhizosphere nitrogen cycling plant –microbe interactions root exudates microbial gene regulation plant breeding targets Extracellular vesicles in the rhizosphere: targets to improve nutrient use efficiency of crops? extracellular vesicles; nutrient use efficiency; rhizosphere; nitrogen cycling; plant –microbe interactions; root exudates; microbial gene regulation; plant breeding targets True https://www.frontiersin.org/legal/copyright-statement endstream endobj xref 0 1 0000000248 65535 f 248 2 0000000249 00001 f 0000000000 00001 f 251 1 0000993816 00000 n 381 1 0000994019 00000 n trailer << /Size 382 /Prev 116 /Root 251 0 R /ID [ <0130F331EF8278AA6D0DC8CB513898E8> ] >> %EndExifToolUpdate 993794 startxref 999195 %%EOF 66 0 obj <>/ExtGState<>/Font<>/ProcSet[/PDF/Text/ImageC]/XObject<>>>/Rotate 0/StructParents 0/Type/Page>> endobj 251 0 obj <> endobj 381 0 obj <>stream application/pdf Ivan A. Paponov Stefanie Schulz Michael Schloter Ana Conesa Yehoram Leshem Paponov IA, Schulz S, Schloter M, Conesa A and Leshem Y (2025) Extracellular vesicles in the rhizosphere: targets to improve nutrient use efficiency of crops?. Front. Plant Sci. 16:1681793. doi: 10.3389/fpls.2025.1681793 Copyright © 2025 Paponov, Schulz, Schloter, Conesa and Leshem.. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms. extracellular vesicles nutrient use efficiency rhizosphere nitrogen cycling plant –microbe interactions root exudates microbial gene regulation plant breeding targets Extracellular vesicles in the rhizosphere: targets to improve nutrient use efficiency of crops? extracellular vesicles; nutrient use efficiency; rhizosphere; nitrogen cycling; plant –microbe interactions; root exudates; microbial gene regulation; plant breeding targets True https://www.frontiersin.org/legal/copyright-statement 2025-10-07T13:47:44+08:00 2025-10-07T13:47:44+08:00 2025-10-07T13:47:44+08:00 uuid:740fee56-efd4-4153-9a45-0dad702969c4 uuid:b84ce92a-55df-4555-ac72-04ee62ba496f endstream endobj 382 0 obj <> endobj 383 0 obj <>stream BT /CS0 cs 0.157 0.153 0.161 scn /GS0 gs /T1_0 1 Tf 9 0 0 9 49.6063 752.7117 Tm 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