EFFECT OF HUMIC ACID ON SEED GERMINATION OF SUB-TROPICAL HALOPHYTES UNDER SALT STRESS

Transcription

1 Pk. J. Bot., 49(6): 79-88, 17. EFFECT OF HUMIC ACID ON SEED GERMINATION OF SUB-TROPICAL HALOPHYTES UNDER SALT STRESS SONIA BANO ABBASI 1, BILQUEES GUL 1*, NASIRUDDIN KHAN 2, SALMAN GULZAR 1, AND MUHAMMAD ZAHEER AHMED 1 1 Institute of Sustinle Hlophyte Utiliztion, University of Krhi, Krhi-75270, Pkistn 2 Deprtment of Chemistry, University of Krhi, Krhi-75270, Pkistn * Corresponding uthor s emil: ilqueesgul@uok.edu.pk; Phone: (9221) ; Fx: (9221) Astrt This study ws rried out to determine the effets of exogenous pplitions of three humi id types (HA1,, ) nd their onentrtions (250 nd 50 ) on seed germintion of ten hlophytes. The speies inluded Aeluropus lgopoides, Arthronemum mrosthyum, Cyperus onglomertus, Desmosthy ipinnt, Hlonemum stroileum, Hlopeplis perfolit, Hlopyrum murontum, Phrgmites krk, Sporoolus ioldos nd Urohondr setulos wheres, tretments used were (1) thermoperiod (10/ nd /30 o C); (2) photoperiod (12/12h, light/drk; nd 24h drk) under nonsline onditions. Effets of humi ids on llevition of seed germintion were lso studied in response to 1) the individul nd omined effets of NCl tretments (reported to use 50% germintion inhiition in speies tested) nd 2) photoperiod (12/12h, light/drk; nd 24h drk). In non-sline onditions, humi id did not llevite seed germintion under the su-optiml thermoperiod (10/ o C) in most hlophyte seeds studied exept for tht of HA1 in A. lgopoides (34%), A. mrosthyum (15%), D. ipinnt (12%) nd H. perfolit (%). Humi id did not improve seed germintion in either photoperiod under non-sline onditions t the optiml thermoperiod (/30 o C). In the presene of the light (12/12h; light/drk photoperiod), humi id llevited seed germintion of U. setulos under sline onditions ut not in ny of the other test speies. Under sline ondition nd omplete drkness, the lower onentrtion tretment (250 mg L - 1 ) of ll three humi id types signifintly improved seed germintion of A. lgopoides, C. onglomertus, D. ipinnt, H. perfolit, P. krk nd U. setulos. Under sline onditions nd sene of light, higher onentrtion (t 50 ) of ll humi id types llevited seed germintion in S. ioldos however, no effet of either humi id type ws oserved on seed germintion of H. stroileum. In onlusion, HA ould prtilly sustitute the light requirement for seed germintion of test speies under sline onditions. Key words: Hlophytes, Humi id, Light, Seed germintion, Temperture. Introdution Aout 7% of the world s totl lnd re is ffeted y slinity. In Pkistn, more thn 6.3 million h of griulturl lnd hs eome sline (Yensen, 1995; Flowers & Colmer, 08). This hs led to derese in rop yield espeilly in rid nd semi-rid regions (Yensen, 1995; Tester & Dvenport, 03; Ashrf & Foold, 07; Flowers & Colmer, 08). Hlophytes re nturlly resistnt to high soil slinity nd pper to hve n oligte slt requirement for optiml growth however, most re sensitive to sline environment st seed germintion (Bewley & Blk, 1982). Most hlophytes n resist onsiderly higher levels of slinity ompred to slt sensitive rops t the germintion level. Osmoti nd/or ioni effets of slinity on seed germintion hve een studied t physiologil, iohemil nd moleulr levels (Mhjn & Tutej, 05; Munns & Tester, 08). Hlophyte seed germintion is usully higher in the sene of slinity (Khn et l., 01; Bu et l., 08; Gun et l., 09). Considerle vrition in slt resistne t germintion hs een reported for some hlophytes from low (Hlopeplis perfolit, Mhmoud et l., 1983), to sustntilly high levels (Sliorni here,1700 mm NCl).Seed germintion of Sroorni frutios, Arthronemum mrosthyum nd Hlonemum stroileum ws inhiited y osmoti rther thn ioni effets under sline onditions (Pujol et l., 00). Wheres, inhiition of seed germintion in Aristid dsensionis, Artemesi ordosi (Toe et l., 1999) nd Prosopis stromulifer (Sos et l., 05) ws used y ion toxiity. Inhiition of seed germintion ould our due to disturnes t vrious stges suh s imiition, ell-wll loosening, moiliztion of food reserves nd rdile elongtion (Khjeh-Hosseini et l., 03; Ky et l., 06). Other environmentl extremes ffeting seed germintion inlude light nd temperture (Zehr et l., 13). Phytohromes re reported to e involved in light responses of photolsti seed during germintion (Fenner & Thompson, 05). Hlophytes vry in their optiml light nd temperture requirement for seed germintion (Gul et l., 13).Su-optiml tempertures lso inhiited seed germintion possily y ltering ion flux nd enzyme kinetis (Tlig et l., 08). Most sutropil hlophytes germinte est t moderte (/30 o C; night/dy) thermoperiods while ooler (10/ o C) temperture regimes re inhiitory for seed germintion (Khn & Gul, 02; Gul et l., 13). Exogenous pplition of hemils ould llevite seed germintion of hlophytes under su-optiml photoperiod nd thermoperiod nd slinity regimes (Gulzr & Khn 01; Msindro et l., 02; Brtels & Sunkr, 05; Ymguhi & Blumwld, 05). Applition of GA3, kinetin, thioure, nitrte, ethephone, fusioin, proline, nd etine llevited seed germintion of

2 SONIA BANO ABBASI ET AL., hlophytes (Gulzr & Khn, 01; Ahmed et l., 14). Humi id ould lso e used to improve seed germintion nd improve our understnding of underlying mehnisms of slt resistne of hlophytes t this ritil stge of life yle (Khlesro et l., 15). Humi id is the min orgni onstituent of muniipl wste ompost (Senesi et l., 1996).Higher seed germintion due to humi id under sline onditions is ttriuted to improved soil qulity(higher miroorgnism popultion, nutrient sttus, wter holding pity, orgni ontent (Mdonnell et l., 01). Humi sustnes re previously reported with similr effet to plnt hormones (Nrdi et l., 02; Pizzeghello et l., 13). Applitions of humi id inresed seed germintion of prsley, elery nd leek t vrious temperture regimes (Wlker & Bernl, 04) nd of rley, owpe, whet, en, wtermelon, gernium, uumer nd mrigold seeds under sline nd non-sline onditions (Hrtwigsen & Evns, 00; El-Hefny, 10; Szzepnek & Wilzewski, 11; SilvMtos et l., 12). Litertu reserh etween using the ISI we of knowledge nd using the key words humi id AND hlophyte reveled only 11 reserh rtiles. The present study investigted: 1) the effets of three humi ids on seed germintion of ten su-tropil hlophytes in response to vrious thermoperiod nd photoperiod regimes with nd without NCl tretments. Mterils nd Methods Seed olletion nd storge: Seeds of ten hlophyti speies (Aeluropus lgopoides, Arthronemum mrosthyum, Cyperus onglomertus, Desmosthy ipinnt, Hlonemum stroileum, Hlopeplis perfolit, Hlopyrum murontum, Phrgmites krk, Sporoolus ioldos nd Urohondr setulos), were olleted from their popultions. The list of test speies nd experimentl onditions used in this study re given in Tle 1. Seeds were dethed from infloresene, surfe sterilized with 0.85% Clorox (ommeril NOCl), ir dried nd stored in plsti ottle t 4 o C. Extrtion of humi id (HA) from ol: Col smples were thoroughly wshed, oven dried for 24 h t 105 C nd ground to mesh size #. Humi id ws extrted from ol (10g) dded to 50 ml of 0.25 M NOH nd llowed to mix on n oritl shker for 24 h. The smples were entrifuged t 0 r.p.m. nd filtered thrie to seprte from residue. The filtrte ws idified to ph 1 with the help 6 M HCl nd preipittes llowed to settle overnight efore entrifugtion to reover the humi id rystls. The rystls of humi id were wshed with ml of 0.5% hydrohlori-hydrofluori ids (HCL-HF) solution nd then repetedly rinsed with distilled wter to eliminte inorgni impurities. After drying the purified HA for 24h t 105 C the yield of HA ws lulted. Seed germintion tests: All germintion experiments were rried out in 50 9 mm, ir-tight, plsti Petri pltes. Seeds were treted with three humi id types; HA1 nd (derived from mined ol) nd ommerilly ville (Sigm Aldrih G) form of humi id. Four replites of twenty-five seeds eh filled with 5 ml of respetive test solution were used in this study. Experiment-1 Effet of humi id nd thermoperiod on seed germintion of hlophytes under non-sline onditions: Seeds of eh of the ten ove-mentioned hlophyte speies, were pled in progrmmed inutor st 10/ nd /30 C (12/12h; night/dy) thermoperiod nd llowed to germinte in distilled wter with nd without HA1, nd t 250 nd 50. Seed germintion ws reorded every lternte dy for d. Experiment-2 Effet of humi id nd photoperiod on seed germintion of hlophytes under non-sline onditions: The tretments onsisted of non-sline with three onentrtion (0, 250 nd 50 ) of different types of HA (HA1,, ). The seeds were pled in the Petri pltes nd immersed with test 5ml solutions under progrmmed inutors (Perivl, Boone, USA) with optiml regime (/30 C - night/dy), photoperiod (12/12h, light/drk; nd 24h drk) (25μmol photons m 2 s 1 ; nm Sylvni ool white lmp). Tle 1. List of seleted ten hlophytes with the detil of experimentl onditions. Speies SAL PP TP ( o C) Referene Aeluropus lgopoides 0, 500 L /D 10/; /30 (Gulzr & Khn, 01) Arthronemum mrosthyum 0, 500 L 10/; /30 (Khn, 1999) Cyperus onglomertus 0, L /D 10/; /30 (El-Kelwy et l., 09) Desmosthy ipinnt 0, 500 L /D 10/; /30 (Gulzr et l., 07) Hlonemum stroileum 0, 500 L /D 10/; /30 (Qu et l., 07) Hlopeplis perfolit 0, 500 L /D 10/; /30 (Rsool et l., 16) Hlopyrum murontum 0, 300 L 10/; /30 (Noor & Khn, 1995) Phrgmites krk 0, 500 L/D 10/; /30 (Zehr et l., 13) Sporoolus ioldos 0, 300 L/D 10/; /30 (Gulzr & Khn, 03) Urohondr setulos 0, 0 L /D 10/; /30 (Gulzr et l., 01) Sl-slinity (mm NCl); PP- photoperiod (L: 12 h /12 h light nd drk, D: 24h drk); TP- thermoperiod ( o C) (sed on mentioned referenes)

3 EFFECT OF HUMIC ACID ON SEED GERMINATION UNDER SALT STRESS 81 Tle 2. Anlysis of vrine showing effet of humi id types (HAT) nd onentrtions (HAC) on finl seed germintion of test speies under 12 h photoperiod (drk/light) t different thermoperiods (TP). Speies HAT HAC TP HAT*HAC HAT*TP HAC*TP HAT*HAC*TP A. lgopoides 382 *** 154 *** 989 *** 69 *** 86 *** 36 *** 29 *** A.mrosthyum 138 *** 236 *** 3 ns 102 *** 67 *** 245 *** 56 *** C. onglomertus 4764 *** 2 ns 3724 *** 2 ns 844 *** 10 *** 7 *** D. ipinnt 39 *** 23 *** 266 *** 6 ** 5 ns 17 *** 5 ** H. stroileum 108 *** 11 *** 5 ns 5 ** 50 *** 19 *** 5 ** H. perfolit 2945 *** 17 *** 1764 *** 311 *** 307 *** 2 ns 51 *** H. murontum 15 *** 0 ns 96 *** 5 ** 7 *** 12 *** 7 *** P. krk 16 *** 1 ns 62 *** 13 *** 4 ns 0 ns 5 ** S. ioldos 7 ** 285 *** 651 *** 8 *** 36 *** 341 *** 11 *** U. setulos 6 ** 32 *** 441 *** 17 *** 45 *** 6 ns 1 ns Experiment-3 Effet of humi id nd photoperiod on seed germintion of hlophytes under non-sline nd sline onditions: The tretments onsisted of two NCl onentrtions (non-sline nd inhiitory, onentrtion s mentioned in Tle 1) with three onentrtion (0, 250 nd 50 ) of HA (HA1,, ). The seeds were pled in the petri pltes nd immersed with test solutions under progrmmed inutors with optiml temperture regime (/30 C - night/dy), photoperiod (12/12h, light/drk) (25μmol photons m 2 s 1 ; nm Sylvni ool white lmp). Seeds of test speies (light sensitive seeds Tle 2) were lso germinted in omplete drkness where germintion of seeds kept under 24h drkness ws noted only one fter dys. Seeds were onsidered to e germinted with rdil emergene (Bewley & Blk, 1994). Sttistil nlysis: Germintion dt were rsine trnsformed efore sttistil nlyses to test for homogeneity of vrine. Three-wy nlysis of vrine (ANOVA) were performed with the help of SPSS version 11.0 for Windows (SPSS In., 06) to determine the individul nd intertive effets of vrious prmeters s follows: 1. HA types x HA onentrtions x thermoperiods 2. HA types x HA onentrtions x photoperiods 3. HA types x HA onentrtions x NCl tretments (in 12/12h; drk/light photoperiod) 3. HA types x HA onentrtions x NCl tretments (in 24 drk photoperiod) The post-ho Bonferroni test ws lso rried out to determine differenes etween individul tretment mens. ANOVA nd Bonferroni tests were onduted t p<0.05, on mximum totl of vlues for seed germintion (in 4 replites with 25 su-replites eh). Results Experiment-1 Effet of humi id nd thermoperiod on seed germintion of hlophytes under non-sline onditions: The three-wy nlysis of vrine (ANOVA) indited signifint effets (p<0.05) of HA type nd its intertion with HA onentrtion nd thermoperiod, exept for U. setulos (Tle 2) HA1 signifintly (p<0.0) mitigted inhiitory effet of thermoperiod ompred to un-treted ontrols t 10/ o C on seed germintion of A. lgopoides (34%; HA), A. mrosthyum (15%; 500 mg L - 1 ), D. ipinnt (12%; oth 250 nd 50 ) nd H. perfolit (%), respetively (Supplementry Tle 1) However, HA inhiited seed germintion of C. onglomertus nd P. krk t 10/ o C, while germintion remined unffeted in se of S. ioldos, H. stroileum, H. murontum nd U. setulos (Supplementry Tle 1). Experiment-2 Effet of humi id nd photoperiod (12/12h; light/drk nd 24h drk) under non-sline onditions t /30 o C: The three-wy nlysis of vrine (ANOVA) indited signifint effets of photoperiod (exept for A. lgopoides nd P. krk ) mong ll test speies while there were no intertive effets of ll three prmeters tken together (Tle 3). Two intertions of HA type with HA onentrtion nd of HA type with photoperiod (exept for C. onglomertus) were lso nonsignifint. In generl, humi ids showed no llevition of seed dormny in response to photoperiod under nonsline onditions nd exept for C. onglomertus (in 12/12h; light/drk photoperiod) ompred to un-treted ontrols (Supplementry Tle 2). Experiment-3. Effet of humi id nd slinity tretments on seed germintion in light (12/12h; drk/light) t /30 o C: Three-wy ANOVA of HA type, HA onentrtion, NCl tretments nd their intertions showed signifint (p<0.05) individul nd omined effets on seed germintion of A. lgopoides, C. onglomertus, D. ipinnt, H. perfolit,s. ioldos nd U. setulos (Tle 4). Slinity effets were highly signifint (p<0.001) mong ll test speies s in se of HA type exept for H. murontum, whih remined generlly unffeted y HA type, HA onentrtions nd their intertions (Tle 4). However, HA onentrtion did not ffet seed germintion of A. mrosthyum, H. murontum (Supplementry Tle 2), P. krk nd U. setulos. Among ll speies tested, the sustntil llevitory effet of HA (prtiulrly 50 ) under sline onditions, ws oserved only in U. setulos with either n inhiitory effet or no effet in the other speies (Figs. 1-8).

4 82 SONIA BANO ABBASI ET AL., Tle 3. Anlysis of vrine showing effet of humi id types (HAT) nd onentrtion (HAC) on finl seed germintion of test speies under 12 h photoperiod (drk/light) nd 24 hours drkness. Speies HAT HAC PP HAT*HAC HAT*PP HAC*PP HAT*HAC*PP A. lgopoides 1.6 ns 8.5 *** 1.5 ns 0.3 ns 0.4 ns 5.6 ** 0.3 ns C. onglomertus 15.4 *** 0.5 ns.6 *** 0.7 ns 4.1 * 0.3ns 0.4 ns D. ipinnt 1.6 ns 2.6 * 13.6 *** 0.9 ns 0.2 ns 2.6 * 0.4 ns H. stroileum 1.0 ns 7.9 ** 891 *** 0.6 ns 2.2 ns 1.6 ns 0.2 ns H. perfolit 1.6 ns 11.3 *** *** 0.6 ns 0.1 ns 0.8 ns 0.5 ns P. krk 2.8 * 0.3 ns 22.2 *** 1.7 ns 1.3 ns 0.5 ns 2.1 ns S. ioldos 0.3 ns 12.1 *** 8.8 ** 0.3 ns 0.5 ns 5.5 ** 0.3 ns U. setulos 0.2 ns 1.5 ns 3.5 * 0.7 ns 0.4 ns 0.7 ns 1.2 ns Tle 4. Anlysis of vrine showing effet of humi id types (HAT) nd onentrtions (HAC) on finl seed germintion of test speies under 12 h photoperiod (drk/light) under slinity. Speies HAT HAC SAL HAT*HAC HAT*SAL HAC*SAL HAT*HAC*SAL A. lgopoides 151 *** 146 *** 5596 *** 39 *** 118 *** 48 *** 34 *** A. mrosthyum 7.0 ** 2 NS 98 *** 6 *** 6 *** 9 *** 3 ns C. onglomertus 158 *** 36 *** 2172 *** 38 *** 64 *** 8 *** 6 ** D. ipinnt 10 *** 54 *** 13 *** 7 *** 13 *** 7 ** 5 ** H. stroileum 10 *** 6 *** 191 *** 1 ns 4 ns 1 ns 2 ns H. perfolit 61 *** 4 *** 3472 *** 19 *** 101 *** 71 *** 26 *** H. murontum 1 ns 1 ns 47 *** 2 ns 0 ns 1 ns 1 ns P. krk 23 *** 1 ns 639 *** 19 *** 6 * 4 ns 4 ns S. ioldos 69 *** 1671 *** 5828 *** 61 *** 33 *** 888 *** 19 *** U. setulos 15 *** 3 ns 362 *** 7 *** 33 *** 258 *** 29 *** Tle 5. Anlysis of vrine showing effet of humi id types (HAT) nd onentrtions (HAC) on finl seed germintion of test speies under 24 h drkness under slinity. Speies HAT HAC SAL HAT*HAC HAT*SAL HAC*SAL HAT*HAC*SAL A. lgopoides 36 *** 871 *** 471 *** 11 *** 7 ** 585 *** 5 ** C. onglomertus 1535 *** 12 *** 2249 *** 44 *** 507 *** 62 *** 35 *** D. ipinnt 10 *** 18 *** 28 *** 4 ns 2 ns 11 *** 3 ns H. stroileum 2 ns 111 *** 55 *** 5 ** 1 ns 37 *** 0 ns H. perfolit 26 *** 199 *** 290 *** 31 *** 1 *** 163 *** 5 *** P. krk 52 *** 15 *** 517 *** 49 *** 22 *** 59 *** 25 *** S. ioldos 24 *** 296 *** 113 *** 0 ns 17 *** 1059 *** 1 ns U. setulos 0 ns 18 *** 2 *** 14 *** 28 *** 22 *** 4 ns Experiment-3. Effet of humi id nd slinity tretments on seed germintion in drk (24h omplete drkness) t /30 o C: Three wy ANOVA indited signifint effets of HA onentrtion nd slinity tretments nd their intertion on seed germintion of ll hlophyte test speies under omplete drkness (exept for D. ipinnt, S. ioldos nd U. setulos) (Tle 5). In generl, ll HA types ( ) enhned seed germintion of A. lgopoides (%) (Fig. 1), C. onglomertus (%) (Fig. 2), D. ipinnt (%) (Fig. 3), P. krk (%) (Fig. 6), H. perfolit (10%) (Fig. 5) nd U. setulos (%) (Fig. 8), exept of S. ioldos (%) (Figure 7) where high onentrtion of HA (50 ) ws more effetive in omprison with untreted seeds for eh speies in omplete drk long with slinity. However, HA showed no effet on seed germintion of H. stroileum (Fig. 5). Disussion Humi id ws reported to improve seed germintion of mny ultivted speies (Ser nd Novk, 11; Turkmen et l., 04; Antosov et l., 07; Nmdrn-Goorn et l., 14; Akini, 09; Ptil & Wdje, 10; Msindro et l., 07; Erhimi & Miri, 16). However, inhiitory effets of HA were lso reported for germintion of rley seeds (Cvusoglu et l., 15), with no effet in lettue nd tomto (Piolo et l., 1993; Turkmen et l., 04; Antosov et l., 07).

5 EFFECT OF HUMIC ACID ON SEED GERMINATION UNDER SALT STRESS 83 Tle 6. Summry of the effets of humi id (HA1) on seed germintion responses of ten hlophyti speies under su-optiml slinity (50% inhiition), photoperiod (24h drk) nd temperture (10/ o C). Seed germintion Speies Complete drk nd Su-optiml High slinity sline temperture Aeluropus lgopoides Arthronemum mrosthyum ND Cyperus onglomertus Desmosthy ipinnt. Hlonemum stroileum Hlopeplis perfolit Hlopyrum murontum ND Phrgmites krk Sporoolus ioldos Urohondr setulos Arrow indite the signifint inrese (upwrd) or derese (downwrd ompred to their respetive ontrols. Horizontl lines indite no hnge while ND indites speies not use in tht prtiulr experiment Experiment-1 Humi id nd theromoperiod effets: The influene of HA during seed germintion t su-optiml temperture nd photoperiod ws lso determined. HA1 prtilly llevited the effets of the lower temperture regime (10/ o C) on seed germintion of A. lgopoides nd A. mrosthyum wheres, nd mitigted this effet in se of U. setulos nd P. krk. No effet of HA ws found on seed germintion of A. mrosthyum, D. ipinnt, H. stroileum, H. murontum nd S. ioldos t 10/ o C. Humi id is reported to inresed seed germintion of prsley, leek, elery, tomto, lettue, sil, rdish nd grden ress vegetle speies in wide rnge of tempertures (Yildirim et l., 02). Su-optiml tempertures either dely seed germintion or indue seed dormny y ltering hormonl lne. The kinetin-like hormonl property of HA ppers to overome dmging effets of su-optiml tempertures on seed germintion s in se of sugr eet (Akeson et l., 19). Seed germintion inhiition t low temperture ws ssoited with prolems in inding of nitrte with nd tivtion of phytohromes involved in GA prodution (Kendrik & Kronenerg, 12). HA possily inreses the nitrogen ontents of seeds there y skipping the role of phytohrome reeptors (Keuskmp et l., 10; Bouwmeester & Krssen, 1989) s lso reported in Spergul rvensis (Krssen et l., 1988). Experiment 2 - Effets of humi id nd photoperiod: Humi id ppered to inhiit seed germintion of ll test speies under oth thermoperiods tested possily due to the presene of phenoli ompounds (Pellisier, 1993; Hlley & Pellisier, 1997) s in se of some onifers nd leguminous speies (Musolo et l., 13). Another possile reson ould e the derese in wter potentil s result of high wter inding ility of humi id (Pettit et l., 04). Experiment-3 Effets of humi id, photoperiod nd slinity in response to light (12/12h; light/drk): Applition of HA improved seed germintion of grden ress, whet, prsley, elery nd leek ws under NCl tretments (Yildirim et l., 13; Msindro et l., 02; Al-Erwy et l., 16), ut not in se of onion (Yildirim et l., 14). On the ontrry, inhiitory effets of HA were reorded in Droephlum moldvi nd Sturej hortenssis (Khlesro et l., 15). In the present study, humi id improved the finl germintion of U. setulos y 2 folds under sline onditions prtiulrly in the higher HA onentrtion (50 ) tested. Effets of HA ppered to e relted to ROS signling (Billy et l., 08) for rdile emergene during seed germintion (Cordeiro et l., 11). In ddition, llevitory effets of HA on proesses suh s respirtion, ell division, minerls uptke on seed germintion under sline onditions, indite hormone-like tivity (Pettit, 04). HA enhned the ethylene prodution under sline onditions vi the ETHYLENE RESPONSE FACTOR1 (Mor et l., 12).

6 84 SONIA BANO ABBASI ET AL., Germintion (%) HA1 ) Light ) Drk NCl (mm) 50 Germintion (%) HA1 ) Light ) Drk NCl (mm) 50 Fig. 1. Effet of humi id types (HA1,, ) nd HA onentrtions (0, 250, 50 ) on seed germintion of Aeluropus lgopoides under non-sline nd sline onditions (ffeting 50% germintion inhiition, reported for eh speies) t photoperiod )12/12h; light/drk, ) 24h drk; /30 o C. Brs indite mens (± SE; n = 4 replites with 25 seeds eh) while different letters over rs indite signifint (p<0.05) differenes mong humi id onentrtions (Bonferroni test). Fig. 3. Effet of humi id types (HA1,, ) nd HA onentrtions (0, 250, 50 ) on seed germintion of Desmosthy ipinnt under non-sline nd sline onditions (ffeting 50% germintion inhiition, reported for eh speies) t photoperiod )12/12h; light/drk, ) 24h drk; /30 o C. Brs indite mens (± SE; n = 4 replites with 25 seeds eh) while different letters over rs indite signifint (p<0.05) differenes mong humi id onentrtions (Bonferroni test). Germintion (%) HA1 ) Light ) Drk NCl (mm) Germintion (%) ) Light ) Drk HA NCl (mm) Fig. 2. Effet of humi id types (HA1,, ) nd HA onentrtions (0, 250, 50 ) on seed germintion of Cyperus onglomertus under non-sline nd sline onditions (ffeting 50% germintion inhiition, reported for eh speies) t photoperiod )12/12h; light/drk, ) 24h drk; /30 o C. Brs indite mens (± SE; n = 4 replites with 25 seeds eh) while different letters over rs indite signifint (p<0.05) differenes mong humi id onentrtions (Bonferroni test). Fig. 4. Effet of humi id types (HA1,, ) nd HA onentrtions (0, 250, 50 ) on seed germintion of Hlonemum stroileum under non-sline nd sline onditions (ffeting 50% germintion inhiition, reported for eh speies) t photoperiod )12/12h; light/drk, ) 24h drk; /30 o C. Brs indite mens (±SE; n = 4 replites with 25 seeds eh) while different letters over rs indite signifint (p<0.05) differenes mong humi id onentrtions (Bonferroni test).

7 EFFECT OF HUMIC ACID ON SEED GERMINATION UNDER SALT STRESS 85 Germintion (%) HA1 ) Light ) Drk NCl (mm) Germintion (%) HA1 ) Light ) Drk NCl (mm) Fig. 5. Effet of humi id types (HA1,, ) nd HA onentrtions (0, 250, 50 ) on seed germintion of Hlopeplis perfolit under nonsline nd sline onditions (ffeting 50% germintion inhiition, reported for eh speies) t photoperiod )12/12h; light/drk, ) 24h drk; /30 o C. Brs indite mens (± SE; n = 4 replites with 25 seeds eh) while different letters over rs indite signifint (p<0.05) differenes mong humi id onentrtions (Bonferroni test). Germintion (%) HA1 ) Light ) Drk NCl (mm) 50 Fig. 6. Effet of humi id types (HA1,, ) nd HA onentrtions (0, 250, 50 ) on seed germintion of Phrgmites krk under nonsline nd sline onditions (ffeting 50% germintion inhiition, reported for eh speies) t photoperiod )12/12h; light/drk, ) 24h drk; /30 o C. Brs indite mens (±SE; n = 4 replites with 25 seeds eh) while different letters over rs indite signifint (p<0.05) differenes mong humi id onentrtions (Bonferroni test Fig. 7. Effet of humi id types (HA1,, ) nd HA onentrtions (0, 250, 50 ) on seed germintion of Sporoolus ioldosunder nonsline nd sline onditions (ffeting 50% germintion inhiition, reported for eh speies) t photoperiod )12/12h; light/drk, ) 24h drk; /30 o C. Brs indite mens (± SE; n = 4 replites with 25 seeds eh) while different letters over rs indite signifint p<0.05) differenes mong humi id onentrtions (Bonferroni test) Germintion (%) HA1 ) Light ) Drk NCl (mm) 50 Fig. 8. Effet of humi id types (HA1,, ) nd HA onentrtions (0, 250, 50 ) on seed germintion of Urohondr setulos under nonsline nd sline onditions (ffeting 50% germintion inhiition, reported for eh speies) t photoperiod ) 12/12h; light/drk, ) 24h drk; /30 o C. Brs indite mens (± SE; n = 4 replites with 25 seeds eh) while different letters over rs indite signifint (p<0.05) differenes mong humi id onentrtions (Bonferroni test).

8 86 SONIA BANO ABBASI ET AL., Supplementry Tle 1. Effets of humi id type (HA1,, ) nd HA onentrtion (0, 250, 50 ) on seed germintion of ten hlophyti speies t different thermoperiod (TP; 10/ nd /30 o C). Speies TP ( o C) C HA A. lgopoides 10/ 43 ± 1 77 ± 1 52 ± 0 27 ± 1 d ± 0 d 25 ± 1 d ± 0 /30 92 ± 1 94 ± 0 85 ± 4 79 ± 1 65 ± 1 29 ± 0 e ± 0 d A. mrosthyum 10/ 52 ± 0 53 ± 4 64 ± 2 53 ± 1 48 ± 0 52 ± 0 48 ± 0 /30 71 ± 1 45 ± 1 ± 0 51 ± 1 ± 0 d 53 ± 1 d 52 ± 0 C. onglomertus 10/ 0 ± 0 0 ± 0 3 ± 0 0 ± 0 0 ± 0 0 ± 0 15 ± 0 /30 67 ± 1 51 ± 1 48 ± 0 68 ± 0 63 ± 0 68 ± 0 69 ± 1 D. ipinnt 10/ 21 ± 1 33 ± 4 35 ± 3 13 ± 1 ± 2 25 ± 3 ± 2 /30 ± 0 56 ± 4 51 ± 4 39 ± 1 39 ± 1 32 ± 0 28 ± 0 H. stroileum 10/ ± 0 ± 0 ± 0 81 ± 0 81 ± 0 ± 0 ± 0 /30 92 ± 0 85 ± 3 91 ± 5 85 ± 1 ± 0 93 ± 1 ± 0 H. perfolit 10/ ± 0 ± 0 93 ± 1 55 ± 1 d ± 0 d 69 ± 1 52 ± 0 d /30 ± 0 ± 0 ± 0 ± 0 72 ± 0 ± 0 69 ± 1 H. murontum 10/ 88 ± 0 88 ± 2 96 ± 2 89 ± 1 84 ± 4 ± 0 95 ± 1 /30 ± 0 ± 0 ± 0 92 ± 0 96 ± 2 95 ± 1 97 ± 1 P. krk 10/ 47 ± 1 36 ± 0 39 ± 4 49 ± 1 41 ± 0 52 ± 0 52 ± 0 /30 81 ± 1 37 ± 2 41 ± 2 75 ± 1 68 ± 0 71 ± 1 69 ± 0 S. ioldos 10/ 13 ± 1 8 ± 4 11 ± 3 ± 0 16 ± 0 12 ± 0 9 ± 1 /30 61 ±1 13 ± 1 19 ± 1 16 ± 0 8 ± 0 25 ± 1 24 ± 0 U. setulos 10/ 59 ± 1 ± 2 17 ± 1 11 ± 1 8 ± 0 8 ± 0 8 ± 0 /30 85 ± 1 76 ± 6 ± 0 ± 2 25 ± 1 ± 4 30 ± 0 Supplementry Tle 2. Effet of humi id type (HA1,, nd ), HA onentrtion (0, 250, 50 ) nd photoperiod (12/12h; light/drk nd 24h drk) on seed germintion of ten hlophyti speies t /30 o C. Speies pp C HA A. lgopoides L 92 ± 0 94 ± 2 85 ± 7 78 ± 1 65 ± 1 29 ± 1 e ± 0 d D 69 ± 1 65 ± 2 0 ± 0 41 ± 1 0 ± 0 49 ± 1 0 ± 0 C. onglomertus L 67 ± 1 51 ± 1 48 ± 0 68 ± 0 63 ± 1 68 ± 0 68 ± 0 D 53 ± 1 0 ± 0 0 ± 0 45 ± 1 64 ± 0 56 ± 0 43 ± 0 D. ipinnt L ± 0 56 ± 4 51 ± 4 39 ± 1 39 ± 0 32 ± 0 28 ± 0 D 9 ± 1 0 ± 0 0 ± 0 0 ± 0 0 ± 0 0 ± 0 0 ± 0 H. stroileum L 92 ± 0 85 ± 3 91 ± 5 85 ± 1 ± 0 93 ± 1 ± 0 D 24 ± 0 17 ± 3 4 ± 1 12 ± 0 0 ± 0 d 13 ± 1 0 ± 0 d H. perfolit L ± 0 ± 0 ± 0 ± 0 72 ± 0 ± 0 69 ± 1 D 28 ± 0 27 ± 1 27 ± 1 ± 0 0 ± 0 8 ± 0 8 ± 0 P. krk L 81 ± 1 37 ± 2 41 ± 2 75 ± 1 68 ± 0 71 ± 1 69 ± 0 D 44 ± 0 35 ± 3 0 ± 0 21 ± 1 31 ± 1 41 ± 1 37 ± 1 S. ioldos L 61 ± 1 13 ± 1 19 ± 1 16 ± 0 8 ± 0 25 ± 1 24 ± 0 D 25 ± 1 0 ± 0 0 ± 0 7 ± 1 4 ± 0 7 ± 1 4 ± 0 U. setulos L 85 ± 1 76 ± 6 ± 0 ± 2 25 ± 1 ± 4 30 ± 0 D 82 ± 2 72 ± 5 21 ± 2 72 ± 1 15 ± 1 68 ± 0 25 ± 1 Vlues re mens ± SE Supplementry Tle 3. Effet of humi id types (HA1,, ) nd onentrtions (0, 250, 500 mg L 1 ) on seed germintion % A. mrosthyum of nd H. murontum in non-sline (NS) nd sline (S) ondition t 12 h photoperiod with /30 o C. Speies ST C HA A. mrosthyum NS 71 ± 1 45 ± 1 ± 0 51 ± 1 ± 0 53 ± 1 52 ± 0 S 56 ± 0 59 ± 1 45 ± 1 31 ± 3 49 ± 1 52 ± 0 36 ± 0 H. murontum NS ± 0 ± 0 ± 0 92 ± 0 96 ± 2 95 ± 1 97 ± 1 S 29 ± 3 28 ± 3 32 ± 0 37 ± 3 25 ± 1 24 ± 0 24 ± 0 Vlues re mens ± SE

9 EFFECT OF HUMIC ACID ON SEED GERMINATION UNDER SALT STRESS 87 Experiment-3 Effets of humi id, photoperiod nd slinity in response to drkness (24h drk): Under sline onditions, lower dose ( ) of humi id hd positive effet on seed germintion of photolsti seeds exept for H. stroilium, while higher doses (50 ) of humi id hd negtive effets on seed germintion. This dose-speifi response of HA ws lso reported for some other speies (Msindro et l., 02; Erhimi & Miri, 16). The llevition of photolsti seed germintion y humi id from omplete drkness ould e hormone-like tivity (Pizzeghello et l., 01; Sutliffe & Whitehed, 1995). Photolsti seeds hd higher ROS in the presene of light (Leymrie et l., 12). Humi id ould llevite slt indued seed dormny in the sene of light for most of the hlophyti speies tested. Biohemil nd moleulr studies ould further eluidte the underlying mehnisms of these responses. 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