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Journal of Plant Diseases and Protection Zeitschrift für Pflanzenkrankheiten und Pflanzenschutz Sonderheft XX, 471-478 (2006), ISSN 1861-4051 © Eugen Ulmer KG, Stuttgart The allelopathic potential of common dandelion (Taraxacum officinale WEB.) V. GYENES1*, I. BÉRES Institute for Plant Protection, Department of Herbology and Pesticide Chemistry, University of Veszprém, Georgikon Faculty of Agriculture, H-8361 Keszthely, P.O.Box 71. Hungary, e-mail: [email protected] * Corresponding author Summary The common dandelion (Taraxacum officinale WEB.) is one of the most troublesome weeds in meadows, pastures, grasses, turf grasses, orchards and vineyards. The aim of this study was to establish the allelopathic effect of the perennial dicotyledonous weed and to ascertain to what degree is it able to influence the germination of grass species widespread in turfs. Seed of selected Gramineae and Fabaceae species (Lolium perenne L., Arrhenatherum elatior L., Festuca rubra L., Dactylis glomerata L., Poa pratensis L., Lotus corniculatus L., Trifolium repens L., Trifolium fragiferum L., Bromus inermis L., Bromus erectus L.), which are regarded as important components of turfs, were treated with acetonic, ethanolic and aqueous extracts of T. officinale roots and leaves. The effect of treatments was evaluated by determining seed germination of the test species. The acetone and ethanol extracts of dandelion roots generally inhibited seed germination the most. Aqueous, acetone and ethanol extracts of leaves were generally less inhibitive. Germination of test species’ seed decreased as extract concentrations increased. Keywords: Allelopathy, Taraxacum officinale, Gramineae species, Fabaceae species, germination Zusammenfassung Die allelopatische Wirkung von Taraxacum officinale WEB. Mit diesen Versuchen würde überprüft, ob die aus gemahlenen Teilen der Blätter und Wurzeln hergestellten Extrakte die Keimung von Samen hemmen. Die Keimung der Diasporen von 10 Kulturpflanzen wurde durch den Saft von T. officinale in artspecifischer Weise beeinflußt. Die Extrakte aus Pflanzenteilen hatten eine unterschiedliche Wirkung auf die Keimrate von Kulturpflanzensamen. Die Extrakte aus Blättern und Wurzeln von Taraxacum officinale zeigten signifikante Hemmwirkungen auf Blätter (4,87-90,09 %) und auf Wurzeln (0,99-96,56 %). Bei diesem Unkraut wurden in Laborversuchen phytotoxische Wirkungen auf Testpflanzen festgestellt. Stichwörter: Allelopathie, Taraxacum officinale, Grasarten, Kleearten, Keimung Introduction The term ”allelopathy” was first used by MOLISH (1973), and since then it has become generally accepted in the botany and herbology sciences. The word “allelopathy” has Greek origins and stems from the words: ”mutual” and ”influence”. It serves to denote chemical interaction between plants, and therefore, allelopathy means that the donor plant has an inhibitory or stimulatory effect on the acceptor plant. Allelopathy differs from other negative plant interactions, such as competition, because in the former case 472 GYENES, BÉRES the donor plant has a harmful influence on other plants by emission of allelochemical compounds. These compounds are secondary metabolites that often are involved with different protective mechanisms. In Hungary, T. officinale is a widespread weed that is most troublesome in the rainy western part of the country (KARAMÁN 2000). Its importance is demonstrated by data of the Hungarian Weed Survey for vineyards (MIHÁLY 2005). In Sághegy the weed was ranked in 3rd place due to its 5.04 % average cover In Somló it was also ranked in 3rd place based on 2.40 % cover, but in Badacsony it was ranked 5th as a result of 1.41 % cover. The importance of the weed was similar in orchards DANCZA et al. (2005). This is demonstrated by the latest data from the Hungarian Weed Survey for orchards (cherry and apple plantations). Based on average cover % the weed is ranked in 5th place in apple plantations, and in 9th place in apple plantations. The beautiful yellow, shining inflorescences of T. officinale are recognized from a distance in those areas where chemical weed control was not effective. Such localities include orchards, roadsides, gardens, ditches, etc. The weed could become a dominant weed in planted grasses and fodder crops. It is often not found in cereals and row crops under normal crop production conditions (KARAMÁN 2000). Dandelion is a perennial weed with leaves arranged in rosette pattern, and its thick taproot extends vertically to deep soil layers (UJVÁROSI 1973). Its rootstock is reddish-brown and leaf scars of the previous year are visible on it. The rootstock of stronger and older plants is polyandrous. After the death of the principal bud the new shoots develop from lateral buds. In the vegetative stage the different aboveground parts are well developed in a rosette pattern and with a short stem. In the reproductive phase a new scape is raised from the short stem. The capitulum occurs on the end of the scapes (CZIMBERMAGYAR 2005). The capitulum is 2.5-7.5 mm in diameter, and often convex. Inflorescences are 12-25 x 15-25 mm; outer bracts up to 17 mm, linear-lanceolate, usually rather dark, more or less glaucous green, paler on inner surface, pale margin often present but never conspicuous, erect to deflexed, ecorniculate or slightly callosed. Ligules are long, narrow, medium yellow, usually with a brownish stripe (TUTIN et al. 1990). Principal flowering time is middle of April, beginning of May (KARAMÁN 2000). Achenes are brownish, 2.5-3.5 mm, tuberculate or spinulose. The cone is 0.2-0.7 mm long, conical; beak is 7-15 mm, slender (TUTIN et al.1990). The root contains economically-important secondary metabolites, such as: taraxacin, taraxerin, taraxerol, taraxasterol, rubber, sugar, wax, fats, mucilage, choline, and saponin acids (SIMON 1984). Common dandelion is a serious weed because it is able to spread with seed and also vegetatively with its well-developed taproot and leaves arranged in rosette pattern, it can be very harmful in turfgrasses. It is a serious weed in turfgrasses because its optimal germination time in early spring coincides with that of grass seeds. Due to its rapid development it is particularly suppressive towards weakly developed and poorly growing grasses (KARAMÁN 2000). The weed does not tolerate frequent cultivation. Weed numbers can be reduced significantly with welldesigned tillage and stubble-stripping operations. Mechanical control needs to be done continuously in order to reduce plant numbers as well as seed numbers in the seed bank. In small gardens, the size of turfs (lawns) allows for individual dandelion plants weed to be removed by hand. The advantages of this weed control method are that important components of turfs will be not damaged, and weed control will be realized without any herbicides. Disadvantage is that it requires a lot of human energy (KARAMÁN 2000). Where the weed occurs in crops or on roadsides glyphosate or gluphosinate-ammonium herbicides can be used. Gluphosinate-ammonium has a scorching effect therefore it is more effective on weed seedlings. Weed control in turfs is mainly done with auxintype herbicides (KARAMÁN 2000). Materials and methods The test (acceptor) species were: Lolium perenne L., Arrhenatherum elatior L., Festuca rubra L., Dactylis glomerata L., Poa pratensis L., Lotus corniculatus L., Trifolium repens L., Trifolium fragiferum L., Bromus inermis L., and Bromus erectus L. Fresh rootstock and shoots of T. officinale were collected at the flowering stage in Keszthely, from ruderals in June, 2005. Bioassay experiments were set up in Petri-dishes in June, 2005. Fresh roots and leaves of the weed were cleaned with water and cut into small (1-1.5 cm) pieces, which were homo- Allelopathy of Taraxacum officinale 473 genized in a grinder. Based on the technique of Béres (1992), the following treatments were prepared: 50 g fresh leaf/200 ml acetone, 50 g fresh leaf/200 ml ethanol, 50 g fresh root/200 ml water, 50 g fresh root/200 ml acetone, 100 g fresh root/200 ml acetone, 50 g fresh root/200 m ethanol. Soaking of the material in the various solvents lasted 24 hours, and then the mixtures were filtered through filter paper. Not all of the afore-mentioned treatments were tested on every test species. Supernatant volume was placed on filter paper discs in Petri-dishes of 10 cm diameter, and their lids were removed to allow the solvents to evaporate. Thereafter water volume was added to the dry filter paper in the dishes, and 100 seeds were placed on the filter paper in each Petri-dish. Treatments were replicated four times. Allelopathic effect was assessed based on the germination rate of seeds of the various test species. Germination percentage was recorded on 12th, 14th, 21st and 28th day by MSZ 6354/3. The Microsoft Excel statistic programme was used for analyzing the data. Results Certain aqueous, acetone or ethanol extracts of roots or leaves of common dandelion significantly decreased the germination percentage for several Gramineae and Fabaceae species (Tab. 1-10). Tab. 1: The allelopathic effect of Taraxacum officinale on the germination of Lolium perenne’s seeds. Tab. 1: Wirkungen der Extrakte aus Taraxacum officinale auf die Keimung von Lolium perenne Samen. Species Treatments N1 X2 Decrease in germination % 7.82 10.16 2.87 3.65 28.91 30.99 SD3 CV%4 Lolium perenne (perennial ryegrass) Control 4 96 2.828 2.945 50 g fresh leaf/200 ml acetone 4 88.25 5.446 6.153 50 g fresh leaf/200 ml ethanol 4 86.25 4.924 5.708 50 g fresh root/200 ml acetone 4 93.25 2.061 2.210 50 g fresh root/200 ml water 4 92.5 2.516 2.720 100 g fresh root/200 ml acetone 4 68.25 3.403 4.986 100 g fresh root/200 ml ethanol 4 66.25 5.619 8.481 P < 0.01; LSD5% = 5.97 1 N = number of replicates, 2X = mean germination %, 3SD = standard deviation, 4CV% = variation coefficient Tab. 2: The allelopathic effect of Taraxacum officinale on the germination of Arrhenatherum elatius’s seeds. Tab. 2: Wirkungen der Extrakte aus Taraxacum officinale auf die Keimung von Arrhenatherum elatius Samen. Species Treatments N1 X2 Decrease of germination % 48.23 51.78 47.52 80.86 92.91 SD3 CV%4 Arrhenatheru m elatius (tall oatgrass) Control 4 70.5 1.290 1.829 50 g fresh leaf/200 ml acetone 4 36.5 2.362 6.471 50 g fresh leaf/200 ml ethanole 4 34 4.966 1.460 50 g fresh root/200 ml acetone 4 37 3.109 8.402 100 g fresh root/200 ml acetone 4 13.5 1.500 11.111 100 g fresh root/200 ml ethanol 4 5 2.081 41.62 P < 0.01; LSD5% = 4.206 1 N = number of replicates, 2X = mean germination %, 3SD = standard deviation, 4CV% = variation coefficient 474 GYENES, BÉRES Tab. 3: The allelopathic effect of Taraxacum officinale on the germination of Festuca rubra’s seeds. Tab. 3: Wirkungen der Extrakte aus Taraxacum officinale auf die Keimung von Festuca rubra-Samen. Species Treatments N1 X2 Decrease of germination % 76.29 64.75 78.85 74.36 76.93 SD3 CV%4 Festuca rubra (red fescue) Control 4 39 2.160 5.538 50 g fresh leaf/200 ml acetone 4 9.25 3.201 3.460 50 g fresh leaf/200 ml ethanol 4 13.75 2.217 16.123 50 g fresh root/200 ml acetone 4 8.25 1.707 20.690 100 g fresh root/200 ml acetone 4 10 2.106 21.600 100 g fresh root/200 ml ethanol 4 9 4.546 50.511 P < 0.01; LSD5% = 4.203 1 N = number of replicates, 2X = mean germination %, 3SD = standard deviation, 4CV% = variation coefficient Tab. 4: The allelopathic effect of Taraxacum officinale on the germination of Dactylis glomerata’s seeds. Tab. 4: Wirkungen der Extrakte aus Taraxacum officinale auf die Keimung von Dactylis glomerataSamen. Species Treatments N1 X2 Decrease of germination % 24.30 25.36 27.47 79.23 9.51 80.29 SD3 CV%4 Dactylis glomerata (orchardgrass) Control 4 71 3.162 4.453 50 g fresh leaf/200 ml acetone 4 53.75 7.932 14.757 50 g fresh leaf/200 ml ethanol 4 53 2.449 4.620 50 g fresh root/200 ml acetone 4 51.5 8.736 16.963 100 g fresh root/200 ml acetone 4 14.75 4.031 27.328 50 g fresh root/200 ml water 4 64.25 5.057 7.870 100 g fresh root/200 ml ethanol 4 14 4.242 30.300 P < 0.01; LSD5% = 8.153 1 N = number of replicates, 2X = mean germination %, 3SD = standard deviation, 4CV% = variation coefficient Tab. 5: The allelopathic effect of Taraxacum officinale on the germination of Poa pratensis’s seeds. Tab. 5: Wirkungen der Extrakte aus Taraxacum officinale auf die Keimung von Poa pratensis Samen. Species Treatments N1 X2 Decrease of germination % 90.09 82.33 93.54 96.56 1.73 95.69 SD3 CV%4 Poa pratensis (Kentucky bluegrass) Control 4 58 2.943 5.074 50 g fresh leaf/200 ml acetone 4 5.75 7.135 124.086 50 g fresh leaf/200 ml ethanol 4 10.25 5.315 51.853 50 g fresh root/200 ml acetone 4 3.75 0.957 25.52 100 g fresh root/200 ml acetone 4 2 2.160 108 50 g fresh root/200 ml water 4 57 3.651 6.405 100 g fresh root/200 ml ethanol 4 2.5 2.081 83.24 P < 0.01; LSD5% = 5.85 1 N = number of replicates, 2X = mean germination %, 3SD = standard deviation, 4CV% = variation coefficient Allelopathy of Taraxacum officinale 475 Tab. 6: The allelopathic effect of Taraxacum officinale on the germination of Lotus corniculatus’s seeds. Tab. 6: Wirkungen der Extrakte aus Taraxacum officinale auf die Keimung von Lotus corniculatus Samen. Lotus corniculatus (birdfoot deervetch) Species SD3 CV%4 74 70.5 Decrease of germination % 4.87 3.651 2.082 4.933 2.953 57.5 22.3 3.109 5.406 N1 X2 Control 50 g fresh leaf/200 ml ethanol 4 4 100 g fresh root/200 ml ethanol 4 Treatments P < 0.01; LSD5% = 4.82 N = number of replicates, 2X = mean germination %, 3SD = standard deviation, 4CV% = variation coefficient 1 Tab. 7: The allelopathic effect of Taraxacum officinale on the germination of Trifolium repens’s seeds. Tab. 7: Wirkungen der Extrakte aus Taraxacum officinale auf die Keimung von Trifolium repens Samen. Trifolium repens (white clover) Species SD3 CV%4 57.5 52.5 Decrease of germination % 8.69 2.646 2.646 4.601 6.318 41.5 28.83 3.317 6.375 N1 X2 Control 50 g fresh leaf/200 ml ethanol 4 4 100 g fresh root/200 ml ethanol 4 Treatments P < 0.01; LSD5% = 4.82 N = number of replicates, 2X = mean germination %, 3SD = standard deviation, 4CV% = variation coefficient 1 Tab. 8: The allelopathic effect of Taraxacum officinale on the germination of Trifolium fragiferum’s seeds. Tab. 8: Wirkungen der Extrakte aus Taraxacum officinale auf die Keimung von Trifolium fragiferum Samen. Trifolium fragiferum (strawberry clover) Species SD3 CV%4 24.3 23 Decrease of germination % 5.35 1.633 1.258 5.176 7.1 23 5.35 5.477 23.813 N1 X2 Control 50 g fresh leaf/200 ml ethanol 4 4 100 g fresh root/200 ml ethanol 4 Treatments P > 0.05; LSD5% = N = number of replicates, 2X = mean germination %, 3SD = standard deviation, 4CV% = variation coefficient 1 476 GYENES, BÉRES Tab. 9: The allelopathic effect of Taraxacum officinale on the germination of Bromus inermis’s seeds. Tab. 9: Wirkungen der Extrakte aus Taraxacum officinale auf die Keimung von Bromus inermis Samen. Bromus inermis (smooth brome) SD3 CV%4 81.3 Decrease of germination % - 3.403 4.185 80.5 0.99 3.109 3.862 N1 X2 Control 4 50 g fresh root/200 ml acetone 4 Species Treatments P > 0.05; LSD5% = N = number of replicates, 2X = mean germination %, 3SD = standard deviation, 4CV% = variation coefficient 1 Tab. 10: The allelopathic effect of Taraxacum officinale on the germination of Bromus erectus’s seeds. Tab. 10: Wirkungen der Extrakte aus Taraxacum officinale auf die Keimung von Bromus erectus Samen. N X Control 4 86.3 Decrease of germination % - 50 g fresh root/200 ml acetone 4 71 17.73 Bromus erectus (erect brome) Species Treatments 1 2 SD3 CV%4 2.754 3.191 2.582 3.636 P > 0.01; LSD5% = 4.62 N = number of replicates, 2X = mean germination %, 3SD = standard deviation, 4CV% = variation coefficient 1 From the data in Table 1 it appears that the germination of perennial ryegrass was significantly reduced by both the acetone and ethanol extracts of 100 g root material, by 30.99 and 28.91 %, respectively. Germination of tall oatgrass was influenced similarly as ryegrass (Tab. 2). The germination percentage of this species was 13.5 % in the acetone extract of 100 g root material, and 5 % in the ethanol extract of 100 g root material. Germination of red fescue (Tab. 3.) was significantly reduced by all treatments. The acetone extract of 50 g root material and the ethanol extract of 100 g of the same material had the most pronounced inhibitory effect on the germination of red fescue, with reductions in germination percentage of 78.85 and 76.93 %, respectively. Results for the remaining grass species appear in Tables 4, 5, 9 and 10. From data in Table 4 it appears that the aqueous extract of 50 g root material affected germination of orchard grass the least (64.25 % germination). The aqueous extract was not significantly different from the control treatment, but a significant and similar difference was found between both the acetone and ethanol extracts of 100 g root material and the control treatments (79.23 and 80.29 % germination decrease, respectively). Results for Kentucky bluegrass (Tab. 5.) indicate that germination was significantly (p < 0.01) reduced by all treatments except the aqueous one. Birdfoot deervetch, white clover and strawberry clover are representatives of the Fabaceae family. Results for these species are given in Tables 6, 7 and 8, respectively. For these species only two treatments were applied: 50 g leaf and 100 g root/200 ml acetone. The germination of birdfoot deervetch (Tab. 6) and white clover (Tab. 7) was significantly inhibited by the root treatment. No significant inhibitory effects were observed in the case of strawberry clover (Tab. 8). Discussion Results indicate allelopathic effects for particular acetone, ethanol and aqueous extracts of common dandelion’s leaves or roots on the germination of certain species from the Gramineae and Fabaceae Allelopathy of Taraxacum officinale 477 families. The acetone and ethanol extracts of roots generally had greater inhibitory effects on seed germination than the aqueous extracts. BENDALL (1975) and BÉRES (1992) reported similar results. They pointed out that Cirsium arvense had a significant allelopathic effect, and that root extracts inhibited the germination of crop seeds as well as the development of seedlings. Several articles on allelopathy by Hungarian authors report allelopathic effects. KAZINCZI (2004) examined the allelopathic effect of Cirsium arvense and Asclepias syriaca under bioassay conditions. KAZINCZI (2004) evaluated the allelopathic effect of Ambrosia artemisiifolia, Abutilon theophrasti, Datura stramonium, Cirsium arvense and Asclepas syriaca under laboratory conditions, with sunflower (Helianthus annuus L.) as test plant. She examined the allelopathic effect of extracts of fresh root and leaf material, and the base of evaluation was germination rate. Root extracts had the greatest inhibitory effect on the germination of sunflower. SOLYMOSI (2004) examined the allelopathic effect of root extract of Sorghum bicolor. He pointed out that an aqueous extract of S. bicolor was less inhibitory than an ethanol extract. Own findings suggest that common dandelion is an allelopathic weed shows, theeby emphasizing its importance and significance. With its strong basal development and rosette leaf pattern, it could be very harmful in grasses. It is a serious interferer with planted grasses because in early spring it germinates together with grass seeds. Dandelion’s rapid rate of development gives it a distinct advantage over grasses that do not develop as quickly, or which are weak due to stress conditions. The compounds taraxacin, taraxerin, taraxerol, taraxsterol, which have been found to leach from its roots (SIMON 1984), inhibit the germination and growth of grasses. Acknowledgement The authors whish to thank the Hungarian Scientific Research Found (OTKA) for its financial support (T 037931). References BENDALL, G.M.: The allelopathic activity of Californian thistle (Cirsium arvense (L.) Scop.) in Tasmania. Weed Research. 15, 77-81, 1975. BÉRES, I.: Adatok a Cirsium arvense (L.) Scop. biológiájához. Növényvédelem, XXVII. Évfolyam, 7-8. szám. 322-327, 1992. CZIMBER, GY., L. MAGYAR: Pongyola pitypang vagy gyermekláncfű (Taraxacum officinale WEB.) In: Benécsné Bárdi G. et al. (szerk) Veszélyes 48, veszélyes, nehezen irtható gyomnövények és az ellenük való védekezés. Mezőföldi Agrofórum Kft, Szekszárd, 188-14, 2005. DANCZA, I. Á. TÓTH, L. SZENTEY, G. BENÉCSNÉ BÁRDI, CS. DOMA, L. HÓDI, A. HORNYÁK, CS. KŐRÖSMEZEI, J. MADARÁSZ, F. MOLNÁR, -R. NOVÁK, R. SZABÓ, P. UGHY, L. VARGA: A szőlő- és gyümölcsültetvények első országos gyomfelvételezésének előzetes eredményei. 51.-ik Növényvédelmi Tudományos Napok, Budapest, 2005. KARAMÁN, J.: Szántóföldeink gyomnövényei – Pongyola pitypang (Taraxacum officinale Web.). Agronapló, IV évfolyam, 12, 2000. KAZINCZI, G., I. BÉRES, J. HORVÁTH, A.P. TAKÁCS: Sunflower (Helianthus annuus) as recipient species in allelopathic research. 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