Arch.BioI Sci, Belgrade, 52(4),195-201,2000.

ALLELOPATHY OF PAEONIA OFFICINALISL. 1753 SSP. BANATICA (ROCHEL) s06 1945, A PANNONIAN ENDEMIC AND RELICT SPECIES

L. DJURDJEVICl, ANKA DINICl, VIDA STOJSIC2, MIROSLAVA MITROVIC1, P. PAVLOVIC1 and M. OLDJA3

1Institute for BiologicalResearch Sinisa Stenkovic; 11060Belgrade; 2Institute for Nature Protection ofSerbia, 21000 Novi Sad; 3Porest-Industrial Complex, 13000 Pancevo, Yugoslavia

Abstract - Paeonia officinalis L. 1753 ssp. banatica (Rochel) Soc 1945 represents a Pannonian endemic and rel­ ict plant species. As an endangered, disappearing species, it is protected according to IUCN and included into the Red Book of the Flora of Serbia. In the Deliblato Sands, in a community of English oak (Quercus robur), only a single population of this peony consisting of 74 individuals with a reduced reproduction capacity occurs. Since this could be the consequence of negative allelopathic influence of dominant species of tree, shrub and herbaceous plant layer, we performed allelopathic studies that included qualitative and quantitative analyses of phenolic acids and total phenolics both in the litter and soil of this community. Surface soil layer under Paeonia officinalis L. 1753 ssp. banatica was found to connain 34.36 fig g! and 564.42 fig g! of free and total bound phe­ nols, respectively. A deeper soil layer (10-20 cm) contained much lower amount of free (only 4.44 fig gl) and 571.73 flggl of total bound phenol compounds. In the surface soil layer only three free phenolic acids (p­ coumaric, p-hydroxybenzoic and vanillic acid) in minute amounts (0.57-1.69 fig gl) were detected. In the deeper soil layer p-coumaric acid was absent. Soil covered with Paeonia officinalis ssp. banatica contains five phenolic acids in bound form, p-coumaric and vanillic acid being the most abundant (10.22-30.70 fig gl). These forms are evenly distributed in the surface and deeper soil layer.

UDC 582.675.1 (497.113)

INTRODUCTION

Paeonia officinalis L. 1753 ssp. banatica (RocheI) 1995 the number of generative individuals was dou­ Soc 1945 represents a Pannonian endemic and relict bled and juvenile germinative sprouts appeared plant species. It has a limited range of distribution ex­ (juve-nile 22%, juvenile germinative 49% and gen­ tending to Hungarian Mecsek Mountain, Roumanian erative 30%) (S t 0 j si IS 1995; S t 0 j s i IS et a1 part of the Banat and the Deliblato Sands. In the Deli­ 1995). We hypothesized that such a small number of blato Sands, only one locality - Flamunda and only a individuals and a reduced reproduction capacity single population of this peony in an English oak forest could be the consequence of negative allelopathic influence of dominant plant species from the layer of were referred to in the available literature (C 0 lie trees, shrubs and herbaceous plants. This prompted and B r 0 z 1969). Because of that, this species has been us to perform qualitative and quantitative analyses protected and included into the Red Book of the Flora of phenolic acids and total phenolic compounds both of Serbia. The population consists of a critical number in the litter and the soil under the paeony. Ric e of individuals (74) with a reduced reproduction capac­ (1974) mentioned 15 groups of compounds, most of ity. Until 1994, only about 20% of individuals were gen­ them being secondary metabolites including phe­ erative with an uncertain seed reproduction. In 1994, nolics as significant for allelopathy, due to their in­ the population was transilluminated and as a result, in hibitory action. Because of their universal

195 196 L. DURDEVIC et sl. distribution, high abundance and an important role in med pioneer communities up to complex phytocoeno­ plant life, phenolics were the most extensively studied ses of herbaceous plants and forest trees. Based on the group of compounds interesting from allelopathic point studies of plant cover of this region, sand, steppe, of view. Accumulation of phenolic phytotoxins in soil swamp and forest type of vegetation with the corre­ under dominant plant species within phytocoenoses, sponding associations and subassociations can be leads to the inhibition of seed germination, seedling distinguished (S t j epa nov i c - V e s eli c i c growth, water, phosphorus and calcium uptake, as well 1979). as of photosynthesis. This results in a decreased number Description ofthe bsbitst - The habitat examined of individuals within a community, i.e. complete elimi­ throughout the present study occurs in the central part nation of some plant species (B ate -S mit h 1969; of the Deliblato Sands. It represents a forest clearing L 0 d h i and Ric e 1971; C h 0 u and M u II e r (50 m in diameter) closed from all sides, formed by cle­ 1972; Ric e 1974, 1979; L 0 d h i 1976, 1978; W hit e­ aring of underwood with Rhamnus cstbsrtics as domi­ h cad et a1. 1983; K u i t e r sand Den n e man nant species and individual over 100-year-old trees of 1987; LYu and B I u m 1990; D j u r d j e vic 1991; English oak (Quercus robur). At the edges, old English L i et a1. 1992 a,b; D j u r dj e vic et a1. 1997 a,b). oak (Quercus robur 1.1) and Austrian pine (Pinus njgra 1.1) 18-20 m high trees (30-50 cm in diameter) occur. Tree layer consists of: Populus tremuls 3.3, Populus MATERIAL AND METHODS alba 1.1 and Quercus robur +. Shrub layer is very dense Description of the Deliblato Sands - The Deli­ and includes the following species: Populus tremula 3.3, blato Sands represent a wast, 35 km long and about 20 Populus alba 2.2, Rhamnus cathartka 4.3, Ligustrum km wide region of Southern Banat extending between vulgare 3.3, Viburnum lantana 1.2, Cornus sanguinee the Danube and Carpathian Mountains. Sand masses 2.2, Crataegus monogyna 2.2, Agrimonia eupstoris 2.1, arc forming the dunes in the direction north-west ­ Quercus robur 1.1, Origanum vulgare 1.1, Berberis vul­ south-east with wider or narrower depressions between geris 1.~ Sambucus nigm 1.~ Melilotus otticinale 1.1, them. South-eastern part of the Deliblato Sands is low Erigeron csnsdensis 1.1, Soncbus ervensis +, Cirsium (altitude 70-100 m a.s.l.), while north-eastern one is lanceolatus +, etc. Herbaceous plant layer is composed higher (altitude 193 m a.s.l.). The climate is characteris­ of 51 species including: Erigeron censdensis 2.2, tic for semi-steppe and differs from that in surrounding Rhamnus cethartics 2.~ Calamagrostis cpigeios 3.3, regions. May and June, i.e. the end of vernal period and Clitiopodium vulgare 2.~ Crataegus monogyna 2.~ early summer are characterized by abundant rainfalls. Dactylls glomerata 2.~ Melka trsnsilvsnice 1.~ Ps­ Average water precipitate makes some 633 mm. The coma otticinalis ssp. bsnstica 1.~ Agrimonia eupetoria least amounts of the rainfalls are recorded at the end of 1.1, Rubus caeslus 1.1, Reseda lutes 1.1, Robinia pseu­ summer and beginning of autumn, i.e. during July, August and September. Extreme variations in tempera­ doacada 1.1, Cslsminthe otiicinslis 1.1, Iris vsriegsta ture, not only during the year but also during the day 1.~ Verbascum nigrutn 1.1, Geranium sanguineum 1.1, are the most significant climate characteristic of the Campanula trechelium 1.1, Lotus corniculatus 1.1, Eu­ Deliblato Sands. phorbia cyperississ 1.1, Prunus tenella 1.1, Frsgsris The substrate consisting of eolic drifts represents vesca 1.1, Linsria vulgaris +, Gslium eperine +, Achil­ agenetic or young genetic soil. The substrate made of lea milleiolium +, Potentille tormentills +, Geum ur­ sand of neutral pH is composed of alumino­ banum +, Knsutis sivensis +, Hypericum perioratum ferro silicates, quartz, calcium carbonate and a poor +, Tussilego farfara +, Torilis srvensis +, Melandryum mixture of clay and humous matters. Quartz grains are album +, Lappa major, etc. resistant while alumino-ferrosilicates are susceptible to Phenolic compounds were analyzed in soil sam­ a rapid decomposition. The presence of calcium car­ ples taken from the surface (0-10 cm) and from the bonate acts supporting pedogenetic processes. The soil depth of 10-20 cm, as well as in the samples of partially as a substrate for vegetation is very young and different decomposed litter. After drying, root parts from soil regarding its fertility (D r a k u lie 1969). samples were removed and the samples sieved. Partially In vegetation of the Deliblato Sands, fragments of decomposed litter consisted of leaves and twigs of autochtonous forests and shrubs are succeeded by trembling poplar, hawthorn, white poplar, English oak, groups of sand and meadow-steppe vegetation. In the herbaceous plants, etc.. After drying and grinding, the sand, as a genetically young structureless soil, a gradual samples were sieved (pore size 0.5 mm) and kept in a development of vegetation is expressed from barely for- refrigerator until the analyses. ALLELOPATHY OF PAEONIA OFFIaNAllS L. SSP. BANATICA 197

Analyses ofphenolic compounds a) Analysis of'phenolic compounds In litter - Free forms of phenolics and phenolic acids were extracted from dried litter (4 g) in boiling 80% ethanol and ethyl­ acetate. The bound phenol forms were extracted in ethylacetate after pretreatment procedure involving 60 min boiling of dried material in 2N HCI (M i j d I a et a11975). b) Anelysis ofphenolsin soil - Free forms of phe­ nolics were extracted for 24 h from the samples con­ taining 30 g of dried soil with boiling ethylacetate (3x50 mL). Residual soil was treated with 15 mL of 2N NaOH 0-10 em 10-20 em and after boiling for 24 h the bound phenols were de­ termined (H e nne qui nand Jus t e 1967; Kat a s e Fig. 1. Content of total phenolic compounds in the soil under Psc­ 1 1981 a, b). The quantity of total phenolics (free and onia officinalis ssp. beasties (fig g ). bound forms) in soil samples was detected spectropho­ tometrically, using the Folin-Ciocalteaus reagent (F e ­ I d man and Han k s 1968). The calibration curve was constructed with different concentrations of feru­ lie acid. Qualitative and quantitative analyses of phenolic acids (PA) were performed employing ascending two­ dimensional paper chromatography. This method pro­ 1.4 vides a satisfactory separation of five PA, including two 1.2 1 derivatives of cinnamic acid (ferulic or 4-hydroxy-3 me­ ~g/g thoxy cinnamic acid and p-coumaric or trans-a­ 0.8 hydroxycinnamic acid) and three derivatives of benzoic 0.6 acid (p-hydroxybenzoic acid, vanillic or 4-hydroxy-3­ 0.4 methoxy benzoic acid and syringic or 3,5-dimethoxy-4­ 0.2

hydroxybenzoic acid). These components were separa­ 0-10 em lO-20 em ted using two solvent systems: 1. isopropanol: ethylace­ tate: NH : HzO (30 : 50 : 1 : 19) and 2. 2% acetic 40H Fig. 2. Amount of free phenolic acids in the soil under Paeonia acid. Dried chromatograms were sprayed with p­ officinalis ssp. banatica (figgl). nitro aniline and 20% NaZC03 and developed chroma­ tographic spots were eluted with 45% ethanol. The quantity of PA was determined spectrophotometrically (a Shimadzu UV -160 spectrophotometer), measuring the absorbance of each acid at a characteristic wave­ length (M i j d I a et al. 1975). All analyses were per­ formed in triplicates in three independent experiments.

RESULTS AND DISCUSSION

As shown in Fig. 1, surface soil layer under Pae­ onia offidnaJjs ssp. banstice was found to contain 34.36 ug/g Jig s' free and 564.42 Jig gl total bound phenolic com­ pounds. A deeper soil layer (10-20 em) contained sig­ nificantly lower amount of free phenolics (only 4.44 Jig gl) and 571.73 Jig gl of total bound phenolic com­ pounds, what is almost equal to their concentration in the surface layer. In the soil samples taken from the O-lO em lO-20 em surface layer, only three free phenolic acids (p­ coumaric, p-hydroxybenzoic and vanillic acid) in small Fig. 3. Content of bound forms of phenolic acids in the soil under amounts (0.57-1.69 Jig gl) were detected. Soil sam­ Paeonia officinalis ssp. banatica (figgl) ples taken from the deeper layer contained no p­ coumaric acid (Fig. 2). 198 L. DURDEVIC et al.

Soil under the peony contained five phenolic acids number of generative peony individuals was doubled as bound forms, p-coumaric and vanillic acid being the and juvenile germinative seedlings appeared (S to j sic most abundant (10.22-30.70 J.1g gl). The amount of et al. 1995). However, during the time, sprouts were bound forms of phenolic acids in the surface layer was developed from tree and shrub stumps, and in the stand about equal to that found in the depper soil layer, i.e. with the population of Paeonia officinalis ssp. bsnstica, they were evenly distributed (Fig. 3). K u i t e r sand the process of restoration of English oak phytocoenosis Den n e man (1987) studied phenolic compounds in is going on. Phytocoenological examination shows that podzolic sandy soil under Fagus silvstic« in Nether­ shrub and herbaceous plant layer with over 60 plant lands and found that bound phenolic forms occuring in species is well developed and they can express a nega­ higher amounts (70-80 J.1g gl) than free forms (40-57 J.1g tive effect on the increase of the peony population. The gl) accumulate during autumnal and winter period. results of experimental phytocoenological studies in the These data are in accordance with our earlier results forest of Hungarian oak and Turkey oak on Avala Mt. demonstrating that surface soil layer on Avala Mt. un­ confirmed that Lonicera cspriiolium as a dominant der community Oruo-Ouercetum virgilianae with species in the shrub and herbaceous plant layer expres­ Lithospermum purpureococruleutn as a dominant spe­ sed a negative effect on some herbaceous plants and cies contains 63.22-143.57 J.1g s' of free phenolic forms tree seedlings, as well as on actynomycetes and nitrify­ and 266.67-565.27 J.1g g-l of bound ones (D j u r d j e ­ ing microorganisms (M i sic ct a! 1978, 1979; Din i c vic et al. 1998). et al. 1982, 1983). It has been also reported that the removal of dominant species from shrub and herba­ ceous plant layer leads to an increased number of indi­ viduals and aboveground mass of species previously 450 II p-coumaric • feruiic present in small number and inhibited (C h 0 u and 4(x) o p-hy. benz. []I vanillic Mull e r 1972; Mis i c ct al. 1978, 1979; Din i c et 350 • syringic a! 1982, 1983; Dj u r d j e vic 1989). In this way allelo­ 3(XJ IillI total pathic influence and competition of a dominant species 250 ug/g 2(X) for basic ecological factors within a phytocoenosis gets

150 eliminated and as a result, expansion of a suppressed I(XI and minor plant species occurs.

50 Negative allelopathic influence of trees and shrubs on herbaceous plant species is well documented free bound and it was demonstrated that phenolic compounds play Fig. 4. Amount of free and bound forms of phenolic acids in forest an important role in this process. Phenolic phytotoxins litter (tlggl). occurring in all organs of dominant tree species, especi­ ally in the leaves of Quercus borealis, Q. alba, Platanus In the litter consisting of partially decomposed occidentalis and Celtis occidentalis, express a negative leaves and twigs of trembling poplar, hawthorn, white influence on herbaceous plant species within a phyto­ poplar, English oak, etc. 10.13 mggl of free and 14.09 coenosis. It has been reported that fallen down leaves mggl of total bound phenolic compounds were found. of these tree species contain several phenolic acids, sco­ It contained five phenolic acids with dominance of poline and scopoletine (N a i b and Rice 1971; L 0 d h i bound forms. p-Coumaric, vanillic and syringic acid 1976, 1978). were the most abundant (61.78-194.49 J.1g gl) (Fig. 4). In the community of English oak with the peony Forest litter under different dominant tree species was examined throughout the present study, herbaceous reported to contain five phenolic acids and several phe­ plant layer with 51 species is well developed. Tissues of nolic aldehydes formed by lignin degradation and total different herbaceous species also contain phenolic amount of phenolic compounds ranged from 10.6­ compounds (total phenolics up to 10.98 mg gl) and 20.8 mggl) (K 6 gel and B 0 c h t e r 1985; K 6 g­ free and bound forms of phenolic acids (up to 499.71 e I 1986; D j u r d j e vic and 0 b era n 1995, 1998; J.1g gl) (W hit e h e ad et a! 1983; D j u r d j e vic D j u r d j e vic etal. 1999). et a! 1997a) that can express inhibitory effects on other During 1994 both the trees and the shurbs were plant species (L i eta! 1992 a,b). cleared from the stand. This led to an intensified illu­ Ecological conditions in the Deliblato Sands are mination and a decreased negative allelopathic influ­ rather harsh (frosts, strong winds, drought, deficit of ence of trees and shrubs on the peony. As a result, the soil moisture, shortage of humus and mineral matters). ALLELOPATHY OF PAEONIA OFFICINALIS L. SSP. BANATICA 199

In addition, one should take into account permanent REFERENCES negative effects of anthropological activities on ecosys­ tem as a whole. Under such circumstances, phenolic compounds of trees and shrubs as an additional unfa­ vourable factor can be critical either for elimination or AINaib, F A. G. and Rice, E. L. (1971). Allelopathic effects of Platanus occidentalis. Bull. Torrey Bot. Club 98, 75-82. a decrease in the number of sensitive species individu­ Bate-Smith, E.C (1969). Flavonoid patterns in the monocotyle­ als. This is well documented by a relatively high number dons. In: Perspectives in Phytochemistry. (Eds. Harborne, of protected plant species (C 0 1i c and B r 0 Z 1969; J.B. and Swain, T.), Acad. Press London-New York, Sa in 0 vic 1987). Vegetative period of Paeonis otii­ 167-177. ciJJaJjs ssp. banetice is short and from this point of view Chou, CH and Muller, CH. (1972). 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DaHaTCKII 60lKyp (Paeonia officinaIis L. 1753 ssp. II Ay6JbCM 3CJbllUIHOM cnojy, nOUITO cy paanonepno banatica (Rochel) S06 1945) je naHOHCKII CHAcM II PCJIIIKT. pacnopeheae. Kao yrposceaa spcra xoja anrseaasa, npesta npasanaaa Y CTCJbll xoja ce cacroja OA ACJIIIMll'IHO pa3JIO­ IUCN aaurrahen je II yapmhen Y UpBCHy KIhllry qmopc xceacr JIIIIlIha II rpaH"IIIIW TpCnCTJbllKC, rrrora, 6cnc TO­ Cp611jC. y .D:cJIll6JIaTcKoj Tlenrsapa nocroja casto jCAHa nOJIC, xpacTa nysosaxa II AP. 113MCpCHO je 10,13 Jig gl nonynauaja ca 74 jCAl1HKC Y nysosasosoj sajennana CJIo60AHIIXII 14.09 J1C gl BC3aHllX yxynnax ~CHOJIa. QHa (JIOKaJIIITCT JIaMyHAa) CMaIhCHC pcnpoAyKTIIBHc cno­ caAPlKil nCT ~CHOJIHIIX KIICCJIIIHa, npn "ICMy npeoanahyjy C06HOCTIf, UITO je BcpoBaTHo nocnemma HcraTIIBHor BC3aHII oonana. Hajname asra p-KyMapllHcKC, aaannna­ aJICJIOUaTCKor YTl1~~a AOMIIHaHTHIIX BpcTa 113 cnpaTa CKC II CllPIIHrIIHCKC KIICCJIIIHC (61,78-194,49 pg gl). QHa npseha, lK6yHoBa II 3CJbaCTIIX 611JbaKa aa 6aHaTcKII canpaca 5 ~CHOJIHIIX KllCCJIIIHa, npll "ICMy npeosnahyjy 60lKYp. Crora CMO npacrynarm aJICJIOnaTCKIIM ncrpa­ BC3aHC ~OpMC. HajBIIIlIc llMa p-KyMapllHcKc, BaHIIJIIIHCKC lKIIBaIhIlMa xoja cy ooyxsarana aaanasy II sreperse H CHplIHrHHCKC (61,78-194,49 j.lg gl). .D:IIcKyToBaHa je ~CHOJIHIIX ~cHona KOJIll"IIIHC KIICCJIIIHa II YKynHllx y Moryha yrrora ~CHOJIHIIX jenarsea.a AOMHHaHTHIIX spcra CTCJbll II 3CMJbllUITY OBC sajemrane. PC3YJITaTII noxasyjy 611JbaKa y ~IlTO~CH0311 y cMaIhcIhY 6pOjHOCTH 6olKypa. na nOBpUIIIHCKII CJIoj 3CMJblfUITa non 6aHaTcKIIM 60lKY­ Y nysosaxoaoj 3ajcAHH~1I ca 6aHaTcKHM 60lKY­ 1 pOM, canpacn 34,36 J1C 15 CJI060AHIIX II 564,42 J1C gl pOM, noopo je paaaajen cnpaT lK6yHoBa H 3CJbaCTHX 6H­ ~CHOJIa BC3aHllX yxynaax . .D:y6Jbll cnoj 3CMJblliliTa (10-20 rsaxa (HMa BHIlIC OA 60 611JbHHX npcra), xoje Mory nera­ sm) canpsca 3HaTHO MaIhY KOJIll"IIIHy CJI060AHIIX (casro THBHO YTH~aTII aa noaehaa,e nonynauaje 6olKypa. Kao 4,44 J1C gl) II 571,73 J1C s' yxynnax BC3aHllX ~CHOJIa. y MCpy sa n060JbIlIaIhC YCJIOBa aa oncranax OBC pCJIIIKTHC nOBpIlIIIHCKOM crrojy sescsanrra cy ACTCKTOBaHC casto H CHACMll"IHC BpCTC aanre JIOPC, nOTpc6HO je H3 nenoc­ Tpll CJIo60AHC ~CHOJIHC KIICCJIIIHC (p-KyMapllHcKa, p­ pCAHC 6JIH311HC nonyrrauaje 6aHaTCKor 60lKypa (na pac­ xiiAPOKCll6cH30cBa II BaHllJIIIHcKa) y MaJIIIM KOJIll"IIIHaMa TojaIhy OA 3-4 m) YKJIaIhaTH CBC 6lfJbHC BpCTC xoje aa (0,57-1,69 pggl). .D:y6Jbll CJIoj 3CJbllUITa naje canpscao p­ 611JIO xoja Ha"IHH Mory HcraTHBHO YTIII~aTH (6HOXCMHjCKH xysrapaacxy KIICCJIIIHY. 3CJblliliTC nOA 6aHaTcKIIM 60lKY­ YTH~aj ~CHOJIHHX jenmsea.a AOMHHaHTHIIX BpCTa, pOM caAplKli nCT ~CHonHIIX xacenana y BC3aHOM CTaIhY, 3aCCHa, KOHKYPcH~llja aa BOAY, MHHCpaJIHC MaTcpHjcH npll "ICMy HajBIIIlIc lIMa p-KyMapllHcKC II BaHllJIIIHCKC npocrop). KIICCJIIIHC (10,22-30,70 J1C gl). He nocroja pa3JIIIKa y KOnll"IIIHII BC3aHllX ~CHOJIHIIX KIICCJIIIHa y nOBpIlIIIHCKOM