RESEARCH ON LOW-ALKALOID CONCENTRATION IN VARIETIES OF LUPIN (LUPINUS SP.) IN LITHUANIA. Received: 25 January 2010 Accepted: 25 June 2010

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1 Generl nd Applied Plnt Physiology 2010, Volume 36 (3 4), pp ISSN Pulished y the Institute of Plnt Physiology Bulgrin Acdemy of Sciences Aville online t RESEARCH ON LOW-ALKALOID CONCENTRATION IN VARIETIES OF LUPIN (LUPINUS SP.) IN LITHUANIA Mknickiene Z. nd R. Askviciute * Voke rnch of the Lithunin Reserch Centre for Agriculture nd Forest, Zlioji. 2, LT-02232, Vilnius, Lithuni Received: 25 Jnury 2010 Accepted: 25 June 2010 Summry. Testing for lkloids in lupine vrieties ws conducted t the Voke Brnch of the Lithunin Institute of Agriculture during competitive tril of feeding lupine (Lupinus sp.) in The smples were tken from feeding yellow lupine (Lupinus luteus L), Trkii nd Vilčii vrieties, s well s nrrow-leved lupine (Lupinus ngustifolius L.), Vilnii vriety, nd selection line N1702. Alkloid concentrtion ws estimted in the periods of inflorescence emergence (BBCH 51-55), flowering (BBCH 64-67), development of fruits (BBCH 71-75) nd seed ripening (BBCH 85-88). The test mteril included exsicctes of leves, stems, flowers, pods nd seeds. According to the results, the verge lkloid levels were similr in yellow forge lupine (Lupinus luteus L.) vrieties Trkii nd Vilčii. The Vilnii vriety on verge contined more lkloids thn the nrrow-leved selection line N1702. Anlysis of the verge lkloid levels reveled tht in the yellow lupine (Lupinus luteus L.) vrieties Trkii nd Vilčii lkloid levels in the stems were lower thn in the nrrow-leved (Lupinus ngustifolius L.) vriety Vilnii nd N1702 in ll phеnologicl stges. The verge lkloid level in leves ws lower thn in stems t the stge of flowering (BBCH 64-67). In our study, the highest verge lkloid levels were found in pods (0.114 ± 7) nd flowers (0.114 ± 6) in nrrow-leved lupines, while the lowest level ws found in seeds (2 ± 3) of yellow lupines. Key words: forge lupins; lkloids; vegettion periods; vegettive nd genertive orgns. INTRODUCTION Before 1926, lupines hd een used s sidertes only. E. Buer nd A. Prynishnikov were the first to spek out the nturl existence of lowlkloid lupines, however, reserch into this field ws hmpered y the sence of relile nd rpid methods of determining lkloid plnts (Gtulin, 2002). In 1928, R. Sengush from the Centrl Germn Institute of Genetics proposed method which ws pplied to nlyze 1.5 million of lkloid plnts, nd three non-lkloid mutnts of yellow lupine nd two such mutnts of nrrow-leved lupine were estlished (Kurlovich, 2002). The sence of lkloids ws determined to e n inherited trit, nd the otined individuls y their yields equlled lkloid plnts. *Corresponding uthor: rit.skviciute@voke.lzi.lt

2 Alkloids in lupin 205 These individuls were used in selection work which resulted in the first fmous vrieties of Munchenerg sweet lupines. Low lkloid concentrtion in lupines is derived from iochemicl muttion. The first forge vrieties of lupines were developed y the method of individul selection from lkloid popultions in which low-lkloid mutnts, though rrely, still did occur (Kurlovich, 2002). Alkloid content is dominnt trit which in yellow lupine is determined y four, in nrrow-leved y five nd in white lupine y eight genes (Phn et l., 2007). Crosspollintion of low-lkloid nd lkloid lupine vrieties ws found to produce the lkloid in F 2 nd splitting occurs into lkloid nd non-lkloid genertions t 3:1 rtio. The role of lkloids in plnts is not yet fully cler. Alkloids re supposed to protect plnts from pests which re put off grzing y the cidic tste (Wink nd Hrtmnn, 1982). Another theory proclims lkloids to e useless products of protein metolism (Clements et l., 1996). Yet nother opinion is tht lkloids ccumulted in the underground prts of plnt, prticipte in metolic processes, induce root growth nd, on leching into soil, mke rrier to microorgnisms (Penev, 2006). However, none of the ove theories gives n exhustive explntion of the significnce of lkloids to plnts ecuse some plnts ccumulte them while others do not. Alkloids show n uneven distriution in plnt orgns: some plnts ccumulte them mostly in seeds nd others in leves, roots or cortex, in prenchyml tissue or in cells. The sme plnt my ccumulte oth similr nd different lkloids. During the vegettion period, lkloid concentrtion undergoes chnges, the pek coinciding with the flowering. At the end of vegettion, lkloids ccumulte in seeds nd roots (Hondelmnn, 1984). Alkloid concentrtion in plnt depends on numerous fctors such s ge, environmentl impcts nd geogrphy, lso on how the soil is fertilized (Breitmier, 2002). Lupine (Lupinus sp.) is universl plnt with numerous useful properties. It my e used oth s fodder nd for soil fertiliztion. As fodder, low-lkloid lupine species such s yellow fodder lupine (Lupinus luteus L.) nd nrrow-leved forge lupine (Lupinus ngustifolius L.) re used. Of course, lupines produce lkloids not in order to supply them to mn or nimls. Vrious lkloids function in plnts s insecticides, hericides, fungicides or pest deterrents (Lee et l., 2008; Gtulin, 2002). There is lso n opinion tht lupine lkloids my destroy toxic fungi in forge nd thus fvour forge ssimiltion (Hondelmnn, 1984). There re studies to show tht low levels of lkloids exert no effect on humn nd niml orgnism, while in lrger quntities they my cuse cute ilments or even deth. Lupine lkloids exhiit not only toxic ut lso phrmcologicl properties. In yellow fodder lupine, lkloid concentrtion my vry from 5 to 1.7 while in nrrow-leved lupine from 5 to 3.0. Low lkloid levels in lupines re considered to vry from 5 to Lupine vrieties reeding with low lkloids mount or without lkloids might give new perspectives for lupine use not only in feed production ut lso in the food industry. The im of the present study ws to determine lkloid concentrtion vritions in different lupine vrieties of the species Lupinus

3 206 Mknickiene nd Askviciute luteus L. nd Lupinus ngustifolius L. in the vegettive nd genertive orgns t different developmentl phses. MATERIALS AND METHODS The study ws crried out in t the Voke Brnch of the Lithunin Institute of Agriculture. The experimentl plots were estlished on sndy lom on cronceous fluvil-glcil grvel eluvited soil (IDp), ccording to FAO- UNESCO clssifiction Hplic Luvisols (LVh) with the following grochemicl indices: ph , humus , moile P 2 O 5 nd K 2 O mg kg -1 nd mg kg -1, respectively. Competitive trils of the vrieties were crried out ccording to selection scheme (Mknickiene, 2007). The smples were tken from feeding yellow lupine (Lupinus luteus L), Trkii nd Vilčii vrieties, s well s nrrowleved lupine (Lupinus ngustifolius L.), Vilnii vriety, nd cropper of No Selection line No ws selected y n individul selection method from collection smple No Selected genotype hd low lkloids mount nd intensive pink flower color. Alkloid concentrtion ws estimted in the periods of inflorescence emergence (BBCH 51-55), flowering (BBCH 64-67), development of fruit (BBCH 71-75) nd seed ripening (BBCH 85-88). The test mteril included exsicctes of leves, stems, flowers, pods nd seeds. Freeze-dried plnt mteril ws finely ground t room temperture nd 15 ml 5 (w/v) trichlorocetic cid ws dded to 200 mg plnt mteril. The suspension ws kept t room temperture for 2 h followed y centrifugtion t 3000 rpm for 15 min. An liquot of 12 ml of the superntnt ws susequently lklinized with 25 (v/v) mmoni to ph ~11 nd extrcted twice with 25 ml dichloromethne. The ph ws then rised to 14 y the ddition of 10 M sodium hydroxide nd gin the solution ws extrcted twice with 25 ml dichloromethne. The orgnic extrcts were dried over nhydrous sodium sulphte, collected in flsk contining 100 μg of internl stndrd (n-eicosne) nd concentrted in vcuo. The residues were reconstituted in c. 1 ml ethyl cette. Usully 50 μl phloem sp or 300 μl xylem sp were mde up to 1 ml with 25 (v/v) mmoni. The queous solution ws extrcted four times with dichloromethne (2 ml) nd the comined orgnic extrcts were dried over nhydrous sodium sulphte nd concentrted upon heting (40 C) under continuous strem of nitrogen. The residue ws reconstituted in methnol (100 μl) contining cffeine (10 μg) s internl stndrd. The method of Lee et l. (2007) with modifictions ws used for chromtogrphic nlysis. The lkloid quntities were reclculted s percentge of the dry mtter concentrtion. A sttview ANOVA progrm ws used for sttisticl nlysis of the dt. The otined dt were ssessed y the method of dispersion nlysis, employing the ANOVA (LSD 0.1 ) sttisticl dt processing softwre (Trknovs, 2002). RESULTS AND DISCUSSION In , two forge lupine species (ech with two genotypes) were nlysed for lkloid levels t the stges of inflorescence emergence (BBCH 51-55), flowering (BBCH 64-67), development

4 Alkloids in lupin 207 of fruit (BBCH 71-75) nd seed ripening (BBCH 85-88), seprtely in vegettive nd genertive prts of the plnts. The results showed different lkloid content in different lupine genotypes. The highest lkloid content ws found in leves of Vilnii (0.108 ) t the phse of fruit development (BBCH 71-75) (Tle 1). Anlysis of the verge lkloid levels reveled tht in the yellow lupine vrieties Trkii nd Vilčii lkloid levels in the stems were lower thn in the nrrowleved vrieties Vilnii nd N1702 in ll phenologicl stges. The verge lkloid level in leves ws lower thn in stems t the stge of flowering (BBCH 64-67). The verge lkloid content in leves during inflorescence emergence (BBCH 51-55) ws higher thn in the stems except vriety Trkii. Also, t the stge of flowering (BBCH 64-67) in ll genotypes lkloids in stems were higher thn in leves. At fruit development stge (BBCH 71-75) lkloids in leves were higher thn in stems except for genotype Vilčii. Alkloid levels in lupines undergo distinct periodicl chnges. In plnts, they hve een found to e the intermedite forms of nitrogen metolism, in which these compounds re rendered hrmless nd ccumulte (Kurlovich, 2002; Brchi, 2000). There hs een dt on the possile role of lkloids in the processes of respirtion, oxidtion of vrious compounds such s scoric nd citric cids, hydroquinone, pyrogllol, enzyme synthesis (Lee et l., 2007). The quntittive distriution of lkloids in different stges of lupine development is shown in Fig. 1. The verge lkloid level in cv. Vilnii ws highest t the stges of flowering (BBCH 64-67) (9 ± 3) nd development of fruit (BBCH 71 75) (8 ± 2). Alkloid levels were influenced lso y the species nd genotype. The verge lkloid levels in Tle 1. Alkloid concentrtion in leves nd stems of some lupine species (T. Vokė, , verge dt). Genotype (A) Vegettive orgns (B) Phenologicl growth stges (C), dry mtter Inflorescence emergence (BBCH 51-55) Flowering (BBCH 64-67) Development of fruit (BBCH 71-75) Trkii Vilčii Vilnii Leves ± 9 5 ± ± 3 Stems 6 ± ± 0 6 ± 1 Leves 8 ± ± 2 5 ± 2 Stems 4 ± ± ± 1 Leves 8 ± ± ± Stems 2 ± ± ± N1702 Leves 4 ± 7 2 ± ± Stems ± ± ± LSD 01 (A) = 0.012; LSD 01 (B) = 7; LSD 01 (C) = LSD 01 (AB) = 9; LSD 01 (AC) = LSD 01 (BC) = 9; LSD 01 (ABC) = 7

5 208 Mknickiene nd Askviciute Trkii Vilčii c d c d Vilnii No c d 0 c d Fig. 1. Alkloid concentrtion vritions during different vegettion periods: ) inflorescence emergence (BBCH 51-55), ) flowering (BBCH 64-67), c) development of fruit (BBCH 71-75) nd d) seed ripening (BBCH 85-88) (T. Vokė, , verge dt). the yellow forge lupine (Lupinus luteus L.) vrieties Trkii nd Vilčii were similr. The vriety Vilnii contined on verge more lkloids thn the nrrowleved selection line N1702. In , from the very first developmentl stges, plnts of different vrieties differed in lef color, rnching, growth dynmics. In the flowering phse (BBCH 64-67), the vegettive orgns were finlly formed, nd morphologicl differences mong the vrieties ecme pronounced. We determined lkloid levels in vegettive (leves nd stems) nd genertive (flowers, pods nd seeds) orgns of plnts of four genetic types. The distriution of lkloids in different vegettive nd genertive orgns of lupine plnts is shown in Fig. 2. There re reports tht the sme plnt my contin oth similr nd different lkloids (Kurlovich, 2002; Brchi, 2000). Throughout vegettion, lkloids levels undergo chnges, their pek occurring during flowering. At the end of vegettion, lkloids ccumulte in seedsnd roots (Gtulin 2002, Brummund 1988). In our study, the highest verge lkloid levels were found in pods (0.114 ± 7) nd flowers (0.114 ± 6) in nrrowleved lupines, nd the lowest level ws mesured in the seeds (2 ± 3) of yellow lupines. The min functions of the overground of stem is to develop the lrgest possile re, to sustin the weight of flowers nd fruits nd to intermedite in trnsporting nutrients from roots to leves, flowers nd fruits s well s from leves to roots, flowers nd fruits. Therefore, the stem contins oth conductive nd supportive tissues. Besides, stems often serve s nutritive stores (Crey nd Wink, 1994). Therefore, our study showed tht

6 Alkloids in lupin Leves Stems Flowers Pods Seeds 2 Fig. 2. Alkloid concentrtion in different vegettive nd genertive orgns of lupine: ) yellow lupins (Lupinus luteus L.); ) nrrow-leved lupins (Lupinus ngustifolius L.) (Vokė, , verge dt). lkloid levels in stems were lower thn in pods which re the sic nutritive orgns of plnt. Leves sor CO 2 from the environment nd from roots, vi circultory tissues, receive wter nd minerl slts. Leves, with the id of solr energy, synthesize from this rw mteril vrious orgnic mtters nd supply them lso to the other orgns of plnt (Przyrowski nd Pck, 1994; Breitmier, 2002). Since lkloid concentrtion in plnt constntly chnges throughout 5 the growth period, the mximum stocks of lkloids in leves re ccumulted efore flowering nd they grdully decline together with the qulittive composition of lkloids with respect to the whole lkloid complex (Crey nd Wink, 1994). Alkloid concentrtion in lupines depends on numerous fctors such s species vriety, ge (developmentl stge), environment nd geogrphicl loction. Alkloid concentrtion in plnts hs een found to impct the centrl

7 210 Mknickiene nd Askviciute nervous system of living orgnisms, with low levels cting s stimultors nd higher levels s suppressors. Therefore, the im of lupine selection in Lithuni could e the development of competitive nrrowleved forge lupine vrieties with low lkloid concentrtion. The Voke Brnch of the Lithunin Reserch Centre for Agriculture nd Forest hs ccumulted vlule locl mteril which needs further, more comprehensive selective nd genetic studies. Bsed on the ville ntionl genetic fund of lupines, we could suggest for cultivtion the most suitle lupine specie, suspecies nd vrieties dpted to the Lithunin climtic conditions nd improved in terms of their iochemicl properties (incresed protein concentrtion nd lowered lkloid levels). REFERENCES Brchi S, Luin. Wrschw - Polnd, Breitmier E, Alkloide. Stuttgrt - Germny, Brummund M, Progress in the reeding of yellow lupines. Proceedings of the 5th Interntionl Lupin Conference: Poznn, Polnd, Crey B, M Wink, Elevtionl vrition of quinolizidine lkloid contents in lupine (Lupinus rgenteus) of the Rocky Mountins.J Chem Ecol, 20: Clements JC, BJ Buirchell, WA Cowling, Reltionship etween morphologicl vrition nd geogrphicl origin or selection history in Lupinus pilosus. Plnt Breed, 115: Gtulin GG, Breeding of Lupinus lus cultivrs with different plnt rchitecture. Proceedings of the 10th Interntionl Lupin Conference: Lurvth - Icelnd, Phn HTT, SR Ellwood, K Adhikri, MN Nelson RP. Oliver, The first genetic nd comprtive mp of white lupin (Lupinus lus L.): identifiction of QTLs for nthrcnose resistnce nd flowering time, nd locus for lkloid content. DNA Reserch, Hondelmnn W, The lupin ncient nd modern crop plnt. Theor Appl Genet, 68: 1 9. Kurlovich BS, Lupins. St. Petersurg - Russi, Lee MJ, JS Pte, DJ Hrris, CA Atkins, Synthesis, trnsport nd ccumultion of quinolizidine lkloids in Lupinus lus L. nd L. ngustifolius L. J Exp Bot, 58: Lee ST, KE Pnter, JA Pfister, DR Grdner, KD Welch, The effect of ody condition on serum concentrtions of two tertogenic lkloids (ngyrine nd mmodendrine) from lupines (Lupinus species) tht cuse crooked clf disese. J Anim Sci, 86: Mknickiene Z, Low-lkloid, nrrow-lefed lupine reeding. Zemdiryste-Agricult, 94: Penev A, Stimulting llelopthic effect of plnt extrcts on some crops s fctor for etter germintion nd growth. Proceedings of the 3th interntionl conference on non chemicl crop protection methods: Lille - Frnce,

8 Alkloids in lupin 211 Przyrowski JA, D Pck, Emryo development fter interspecific hyridiztion of Lupinus lus L., Lupinus mutilis Sweet. nd Lupinus ngustifolius L. J Appl Genet, 38: Trknovs P, Dt trnsformtion of iologicl experiments using computer progrm ANOVA. Zemdiryste-Agricult, 77: Wink M, T Hrtmnn, Enzymtic synthesis of quinolizidine lkloid esters: tigloyl-coa: 13-hydroxylupnine O-tigloyltrnsferse from Lupinus lus L. Plnt, 156:

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