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1 Korean J. Plant Res. 27(4): (2014) Print ISSN Online ISSN Original Research Article 종자저장방법및 GA 3 처리가배풍등종자발아와유묘생육에미치는영향 이수광 1, 김효연 1, 구자정 2 * 1 국립수목원유용식물증식센터, 2 국립수목원산림자원보존과 Effects of Seed Storage Methods and GA 3 Application on Seed Germination and Seedling Growth of Solanum lyratum Thunb. Su Gwang Lee 1, Hyo Yun Kim 1 and Ja Jung Ku 2 * 1 Useful Plant Resources Center, Korea National Arboretum, Yangpyeong , Korea 2 Plant Conservation Division, Korea National Arboretum, Pocheon , Korea Abstract - This study was conducted to determine the effects of seed storage methods ( 20, stratification, 2 dry, 2 wet and room temperature) and GA 3 application (control, dh 2O, 10, 100, 1000 ppm) on seed germination and seedling growth of S. lyratum. As a result seed germination rate of S. lyratum was the highest at 91% when seeds were stored at room temperature and then soaked for 24 hours in GA 3 10 ppm. And seedlings of S. lyratum showed the best quality when seed were stored at 2 dry and then soaked for 24 hours in GA ppm, with the growth characteristics of plant height (47 mm ), number of leaves (8), leaf width (12 mm ), leaf length (19 mm ), fresh weight (aerial/root part; 471/476 mg ), dry weight (aerial/root; 106/41 mg ) and seedling quality indices (106). Therefore, S. lyratum seed were stored at 2 dry, and then soaked for 24 hours in GA ppm, seed germination rate was more than 80% and production of superior quality container seedlings. Key words - Solanum lyratum, Seed storage method, Seed germination, Seedling growth, Gibberellin 서언 가지과 (Solanaceae) 가지속 (Solanum) 식물들은주요한식약용식물자원으로감자 (Solanum tuberosum L.), 가지 (S. melongena L.), 까마중 (S. nigrum L.), 배풍등 (S. lyratum Thunb.) 등이이에속한다. 배풍등은줄기의기부만월동하는덩굴성다년초로길이 3 m로자라며울릉도와남부지방에분포한다 (Korea National Arboretum, 2010). 배풍등 ( 排風藤 ) 은풍병을물리치는덩굴이라는의미이며전초를한방에서한약명백모등 ( 白毛藤 ) 이라고도불리며, 청열이습 ( 淸熱利濕 ), 거풍해독 ( 祛風解毒 ) 의효능으로사용된다. 최근에는항종양작용이있어복수암, 자궁경부암억제작용이높고, 신체의항체형성촉진이우수한것으로알려진 (Ahn, 2008) 유용약용자원식물이다. 민간에서는비상초라부르며해열, 해독및황달치료에널 * 교신저자 ( ) : jjku@forest.go.kr 리이용되고있다 (Park et al., 2009). 다양한약효와잠재적효능을지니고있는자원식물의대량증식및생산은이를이용한산업화의첫단계로이미동속식물인감자, 가지, 까마중등은대량증식기술이개발되어 (Oh and Koh, 2012) 산업화까지연계되었다. 지금까지배풍등관련연구로는향균및암세포증식억제효과 (Shin, 2005), 흰쥐에대한간보호효과 (Shin et al., 2003), 화학성분분석및항산화효과 (Shim et al., 1995) 등주로약리학관련연구가대부분이며이는배풍등이자생하는중국과대만에서도마찬가지이다 (Sun et al., 2006; Yang et al., 2012). 이와는반대로대량증식연구는식물조직배양을이용한연구 (Kuo et al., 2012) 가제한적으로진행되었을뿐배풍등의실생번식특히종자저장방법과전처리에따른종자발아와우수묘생산연구는거의진행된바없다. 게다가국내종자발아연구는실내에서진행되는실험이주를이루고이러한결과들이실제포장에서재현되지않는문제 c 본학회지의저작권은 ( 사 ) 한국자원식물학회지에있으며, 이의무단전재나복제를금합니다. This is an Open-Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License ( which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

2 韓資植誌 Korean J. Plant Res. 27(4) : 365~370(2014) 가있다 (Kang and Choi, 2006). 또한가지속식물종자는휴면과낮은발아율로인해유묘출현시기도길고유묘출아율도낮은것으로알려져있다 (Gisbert et al., 2011). 따라서본연구는저장온도와 GA 3 처리에따른배풍등의종자발아와우수묘생산에미치는영향을살펴보고자수행되었다. 재료및방법공시재료본연구에사용된배풍등종자는 2012년 11월 2일제주도서귀포시상효동에자생하는배풍등열매를채종하여육질과종자를분리한다음 2 와상온 (24 ± 4 ) 건조저장하였다. 종자저장방법및 GA 3 처리에따른발아특성종자저장방법은국내자원식물의일반적인저장방법 (Kim et al., 2003) 인 20 저장, 노천매장, 저온 (2 ) 건조및습윤저장, 상온 (24 ± 4 ) 저장의 5가지로하였다. 20 저장및노천매장은 2 건조저장된종자를파종 60일전에저장하였고, 저온습윤저장은파종 30일전에 petridish에여과지 (Whatman No. 1) 2매를깔고종자를넣고증류수 5 ml를공급한후 parafilm 으로밀봉하여저장하였다. GA 3 처리는파종하루전대조구, 증류수, 10, 100, 1000 ppm 의 5가지로달리하여 24시간침지하였다. 처리된종자는 2013 년 4월 18일클라스만 (potgrond H, Klasmann- Deilmann, Germany) 토양이충진된 72구트레이의각구마다한립씩파종하였으며총 24립씩 3반복하였다. 관수는 1일 2회 (08:00 시, 17:00 시 ) 실시하였으며, 모든실험은경기도양평군용문면에위치한국립수목원유용식물증식센터비닐온실에서수행하였다. 자엽이토양을뚫고 2 mm이상돌출된것을발아한것으로기준으로하여, 발아가완료된시점인파종 50일후최종발아율을조사하였다. 종자저장방법및 GA 3 처리에따른묘특성종자저장방법및 GA 3 처리에따른묘생육특성인초장, 엽수 ( 매 ), 엽폭, 엽장, 근장 및생중량 ( mg, 지상 지하부 ) 을각조건당 9개체씩 6월 7일에조사하였으며, 건중량 ( mg, 지상 지하부 ) 은시료를 15일간음지에서충분히자연건조한후전자저울을이용하여측정하였다. 엽폭과엽장은위에서 2번째잎으로완전히전개된것을대상으로측정하였으며, 근장은길이가긴 5개를측정하여평균값을구하였다. 또한묘목품질을나타내는지표 (May, 1985) 로사용되는묘목묘소지수 (SQI; Seedling Quality Indices) 를다음의식을이용하여구하였다. SQI : Total dry weight ( mg ) Plant height Aerial fresh weight ( mg ) + Root length Root fresh weight ( mg ) 통계처리모든통계처리는 SPSS (ver Kor.) 프로그램을사용하였다. 집단간변이분석은이원배치변량분석 (two-way ANOVA) 을통해실시하였으며, 유의성이있는경우 Duncan multiple range test (P = 0.05) 로 2차검증을실시하였다. 결과및고찰종자저장방법및 GA 3 처리에따른발아특성종자저장방법에따른배풍등의발아율은저온건조저장의평균발아율이 86.9% 로가장높았고, 그다음으로상온저장 80.5%, 저온습윤저장 49.4% 순으로나타나 (Table 1) 저온건조저장이배풍등의최적저장방법인것으로판단되었다. 반면, -20 저장종자는평균발아율이 3.3% 로매우저조하였으며, 노천매장저장종자는전혀발아하지않았다. 이처럼저장방법에따라발아율에큰차이를보였으며고도의유의성이인정되었다. 종자저장으로인한휴면은식물이생육하는환경과밀접한관련이있으며이를통해식물재배에대한간접정보를얻을수있다 (Linkies and Leubner-Metzger, 2012). 종자저장온도에서 20 저장과노천매장저장은배풍등이자생하는남부지방의온도와매우다른환경으로깊은휴면에돌입하여발아율이저조한것으로생각된다. GA 3 처리에의해서도배풍등의발아율에차이를나타냈으며, 종자저장방법과마찬가지로고도의유의성이인정되었다. 상온저장의배풍등종자는 GA 3 처리에의해발아율이유의적으로상승하여 GA 3 10 ppm 처리에서최대발아율 91% 를나타내었고, 100 ppm 및 1000 ppm 처리시 83% 와 77% 로나타나 GA 3 농도가높아질수록배풍등의발아율이감소하는경향을보였다. 저온건조저장의배풍등종자는대조구, 증류수및 GA 3 10 ppm 처리에서발아율이각각 88%, 88% 및 90% 로유의적으로동일한수준으로나타났으며 100 ppm 및 1000 ppm 처리시 86% 와 80% 로유의적으로감소하는경향을보였다. 저온습윤저장된배풍등 -366-

3 종자저장방법및 GA 3 처리가배풍등종자발아와유묘생육에미치는영향 Table 1. Effects of seed storage method and GA 3 application on the seed germination rate of S. lyratum GA 3 application (ppm) Con dh 2 O Mean Storage method ± 6.4h z 5.6 ± 6.4h 2.8 ± 2.4h 1.4 ± 2.4h 1.4 ± 2.4h 3.3 Stratification 0.0 ± ± ± ± ± Dry 88.9 ± 6.4ab 88.9 ± 9.6ab 90.3 ± 2.4ab 86.1 ± 8.7bc 80.6 ± 8.7cd Wet 50.0 ± 4.2fg 56.9 ± 13.4f 58.3 ± 4.2f 40.3 ± 2.4g 41.7 ± 8.3g 49.4 Room Temperature 73.6 ± 6.4e 76.4 ± 4.8de 91.7 ± 0.0a 83.3 ± 7.2cd 77.8 ± 8.7de 80.5 Storage method (S) *** GA 3 application (G) ** (S) (G) * z Mean separation within columns by Duncan s multiple range test at P = * Significant at the 5% level. ** Significant at the 1% level. *** Significant at the 0.1% level. 종자에서도 GA 3 처리농도가높아질수록발아율이감소하였다. GA 3 처리는종피에영향을미쳐배발달을도와발아를촉진시키는것으로알려져있으며 (Koornneef, 2002), 가지속식물은 GA 3 처리로인해발아율향상이보고 (Spicer and Dionne, 1961) 된것처럼배풍등또한 GA 3 처리를통해발아율을향상시킬수있었다. GA 3 농도는일반적으로 100 ppm 이사용되며 Solanum stramonifolium 과 S. torvum 에 GA ppm 24시간침지처리시 100% 발아하였다고보고한것 (Hayati et al., 2005) 과달리배풍등은 GA 3 10 ppm 처리에서가장우수하여동속식물이라도 GA 3 처리농도에따라발아율에차이를보이는것으로나타났다. 종자저장방법및 GA 3 처리에따른묘특성배풍등의종자저장방법및 GA 3 처리에따른묘특성을살펴본결과초장은저온건조저장후 GA ppm처리된종자를발아시킨유묘에서 47 mm로나타났으며, 엽수, 엽폭및엽장등의지상부생육특성또한 8매, 12 mm및 19 mm로다른조건에비해가장우수하였다 (Table 2; Fig. 1). 뿐만아니라지상 지하부의생 건중량또한지상 지하부생중량 471 mg / 476 mg및건중량 106 mg / 41 mg으로근장을제외한모든생육특성이가장우수하게나타났다 (Fig. 1, E). 그다음으로초장은저온습윤저장후 GA ppm 처리된종자를발아시킨유묘에서 45 mm로나타났으며, 저온건조저장후 GA ppm 처리된종자를발아시킨유묘에서 40 mm로뒤를이었다. 작물의생육특성중초장은식물의생육특성을살피는기본요소이며 (Yeates et al., 2002), 지상부수량에직접적인영향을주는것으로보고된바있다 (Oh, 2012). 엽수는저온습윤저장후 GA ppm 및 10 ppm 처리된종자를발아시킨유묘에서 7.3매와 7매로저온건조저장후 GA ppm 처리된종자를발아시킨유묘보다 1매정도적게나타났다. 초기유묘생육특성중엽수는후기생육을판단하는지표로사용될수있어 (Lee et al., 2000) 엽수가많은배풍등유묘를정식할시수확량을늘릴수있을것으로기대된다. 엽폭과엽장은저온습윤저장후 GA ppm 및 10 ppm 처리된종자를발아시킨유묘와저온건조저장후 GA ppm 처리된종자를발아시킨유묘에서 9.7 mm이상, 14.9 mm이상의우수한특성을보였다. 지상부생중량은저온건조및습윤저장후 GA ppm 처리된종자를발아시킨유묘에서 283 mg및 250 mg으로나타났으며, 지상 지하부건중량도각각 62 mg / 19 mg및 66 mg / 20 mg으로비슷한경향을나타냈다. 지하부생중량은저온건조저장후 GA 3 처리대조구에서 301 mg으로우수한경향을나타내었으며, 그다음으로저온건조및습윤저장후 GA ppm 처리시 252 mg및 253 mg으로나타났다. 생육특성의요소를종합하여묘목의질을종합적으로판단할수있는묘목묘소지수 (May, 1985) 는근장을제외한모든생육특성이가장우수한저온건조저장후 GA ppm 처리종자의유묘에서 106 으로가장우수하였으며, 그다음조건인저온건조저장후 GA 3 처리대조구의 68보다약 1.5배이상의차이를보였다. 종자저장방법과 GA 3 처리및종자저장방법과 GA 3 처리상호관계에서엽수와근장을제외하고유의성이인정되었다. 엽수는조사시기가생육초기인점으로인해큰차이를보이지않은것으로생각되며, 근장은 72구플러그셀트레이용기의공간이 -367-

4 韓資植誌 Korean J. Plant Res. 27(4) : 365~370(2014) Table 2. Effects of seed storage method and GA 3 application on the seedling quality of S. lyratum Storage method GA 3 Appl. (ppm) Plant height No. of leaves Leaf width Leaf length Root length Fresh weight ( mg /plant) Dry weight ( mg /plant) Aerial * Root * Aerial * Root * SQI (Seedling Quality Indices) 2 Dry Con 31.2ef z de 12.4de d 301.4b 41.1cde 25.9b 68.3b dh 2 O 21.0h f 5.5g gh 54.5i 9.7h 4.0h 10.7d ef bcd 12.4de ef 147.4fg 24.7efg 10.1fgh 31.5bcd bc bc 17.1ab bc 252.8bc 66.0bc 19.2bcd 65.2bc a a 19.2a a 476.2a 106.4a 41.5a 106.4a 2 Wet Con 28.6ef cde 12.2e e 234.7bcd 25.4efg 13.5def 53.1bcd dh 2O 36.8cd cde 12.6cde b 211.2cde 44.0cd 15.4def 41.1bcd de bc 14.9bc bc 237.2bcd 53.8bcd 18.7bcd 51.6bc ab b 15.6b b 253.4bc 62.7bc 20.9bc 53.3bc ef bcd 14.7bc cd 220.2cde 40.2cde 14.8def 45.4bcd Room Temp. Con 27.2fg bcd 12.9cde e 178.1def 28.1def 11.0efgh 41.6bcd dh 2O 27.4fg f 8.7f efg 131.7fg 17.7gh 10.1fgh 34.8bcd gh def 9.7f efg 115.4g 14.7gh 9.8gh 29.2cd h ef 8.0f fgh 120.1g 12.5h 8.3gh 33.1bcd ef de 12.1e e 157.8ef 27.3def 13.1def 42.5bcd Storage method (S) *** ns *** *** ns *** *** *** *** * GA 3 application (G) ** ns *** *** ns *** *** *** ** ** (S) (G) *** ns ** *** ns *** *** *** *** ** z Mean separation within columns by Duncan s multiple range test at P = ns Non significant. * Significant at the 5% level. ** Significant at the 1% level. *** Significant at the 0.1% level. 제한적이기때문에뿌리가충분히발달될수없었던 (Shin, 2009) 것으로생각된다. 종자에 GA 3 처리시유묘생육이촉진되는사례는 Cichorium endivia 과 C. intybus (Tzortzakis 2009), 고추냉이 (Wasabia japonica; Choi and Lee, 1995), 금어초 (Antirrhinum majus; Kang and Choi, 2006) 등에서보고되었으며, 주요이유는 GA 3 처리로발아기간이단축되면서다른조건에비해생육기간이길어초기생육이상대적으로월등했기때문 (Park et al., 2013) 인것으로생각된다. 하지만 GA 3 처리시비이상적인웃자람으로유묘품질에부정적영향을미칠수있으나본연구에서는비이상적인생장을관측할수없었고, 오히려전반적인생육특성이 우수하였다. GA 3 처리대조구에비해증류수처리시묘목묘소지수는저온건조저장 58, 저온습윤저장 12 및상온저장 7 이상의차이를보여증류수처리는배풍등생육에긍정적인영향을미치지않는것으로판단된다. 이는동속식물인까마중종자의수분처리시생장이촉진된다는보고와상이하였으며 (Oh, 2012), 종에따라다른결과를나타내는것으로사료된다. 이상의결과로부터저온건조저장후 GA ppm 처리된종자를파종하면가장우수한배풍등유묘를확보할수있는것으로판단된다

종자저장방법및 GA 3 처리가배풍등종자발아와유묘생육에미치는영향 Fig. 1. Effects of seed storage method and GA 3 application on the seedling quality of S. lyratum. A, 2 Dry GA 3 con. B, 2 Dry dh 2O. C, 2 Dry GA 3 10 ppm.

5 종자저장방법및 GA 3 처리가배풍등종자발아와유묘생육에미치는영향 Fig. 1. Effects of seed storage method and GA 3 application on the seedling quality of S. lyratum. A, 2 Dry GA 3 con. B, 2 Dry dh 2O. C, 2 Dry GA 3 10 ppm. D, 2 Dry GA ppm. E, 2 Dry GA ppm. F, 2 Wet GA 3 con. G, 2 Wet dh 2O. H, 2 Wet GA 3 10 ppm. I 2 Wet GA ppm. J, 2 Wet GA ppm. K, Room temperature (RT) GA 3 con. L, RT dh 2 O. M, RT GA 3 10 ppm. N, RT GA ppm. O, RT GA ppm. 적요본연구는유용약용자원식물배풍등의대량생산을위한기초연구로종자저장방법과 GA 3 처리가종자발아와묘생육특성에미치는영향을구명하고자수행되었다. 종자저장방법은 20 저장, 노천매장, 저온 (2 ) 건조및습윤저장, 상온 (24±4 ) 저장의 5가지로하였으며, GA 3 처리는대조구, 증류수, 10, 100, 1000 ppm 의 5가지로하였다. 그결과배풍등종자를상온저장한후 GA 3 10 ppm 24 시간처리하였을때 91% 의최대발아율을나타내었다. 배풍등의묘특성은저온건조저장후 GA ppm 처리된종자를발아시켰을때초장 47 mm, 엽수 8매, 엽폭 12 mm, 엽장 19 mm, 생중량 ( 지상 / 지하부 ) 471 mg / 476 mg, 건중량 ( 지상 / 지하부 ) 106 mg / 41 mg및묘목묘소지수 106 으로우수묘를확보할수있었다. 따라서배풍등종자는저온저장후 GA ppm 처리시 80% 이상의발아율과양질의우수묘확보가가능한것으로판단된다. References Ahn, D.K Illustrated Book of Korean Medicinal Herbs. Kyohaksa, Seoul, Korea. p. 78 (in Korean). Choi, S.Y. and K.S. Lee Effect of plant growth regulators on the germination and seedling growth of Wasabia japonica Matsum seeds. Korean J. Medical Crop Sci. 3: (in Korean). Gisbert, C., J. Prohens and F. Nuez Treatments for improving seed germination in eggplant and related species. Acta Horticulturae 898: Hayati, N.E., S. Sutevee and J. Sunanta Seed germination enhancement in Solanum stramonifolium and Solanum torvum. Natl. Sci. 39: Kang, J.S. and I.S. Choi Effect of plant growth regulators and seed priming treatment on the germination and early growth of snapdragon (Antirrhinum majus). Kor. J. Life Sci. 16: (in Korean). Kim, S.H., C.H. Lee, H.G. Chung, Y.S. Jang and H.S. Park The germination characteristics of seed in Kalopanax septemlobus Koidz. by storage methods and GA 3 concentrations. J. Kor. For. Soc. 92: (in Korean). Koornneef, M Seed dormancy and germination. Curr Opin Plant Biol. 5: Korea National Arboretum A Field Guide to Trees & -369-

6 韓資植誌 Korean J. Plant Res. 27(4) : 365~370(2014) Shrubs. Geobook, Seoul, Korea. p. 648 (in Korean). Kuo, C.I., C.H. Chao and M.K. Lu Effects of auxins on the production of steroidal alkaloids in rapidly proliferating tissue and cell cultures of Solanum lyratum. Phytochem Anal. 23: Linkies, A and G. Leubner-Metzger Beyond gibberellins and abscisic acid: how ethylene and jasmonates control seed germination. Plant Cell Rep. 31: Lee, Y.H., J.M. Park, S.T. Lee, D.S. Chung and H.K. Kim Effect of seed treatments on germination and growth of Agrimonia pilosa Ledeb. Korean J. Medical Crop Sci. 8: (in Korean). May, J.T Chapter 9; Seedling Quality, Grading, Culling and Counting. Southern Pine Nursery Handbook. Atlanta, USA. p. 3. Oh, J.S Seed germination and plant growth characteristics of Solanum nigrum Linne influenced by seed soaking and chilling treatments. MS Thesis, Sunchon National Univ., Suncheon, Korea. p. 17, p. 21 (in Korean). Oh, S.J. and S.C. Koh Adventitious shoot formation and plant regeneration from explants of Solanum nigrum L. Korean J. Plant Res. 25: (in Korean). Park, J.H., S.S. Park and J.Y. Bae Pharmacognostical studies on the folk medicine Bae Pung Dung. Kor. J. Pharmacogn. 40:25-31 (in Korean). Park, J.S., Y.I. Choi, Y.H. Kim, S.H. Lee, K.N. Kim, M.C. Suh, G.J. Kim and G.J. Lee Optimum germination temperature and seedling root growth characteristics of Camelina. CNU J. Agricult. Sci. 40: (in Korean). Shim, K.H., H.S. Young, T.W. Lee and J.S. Choi Studies on the chemical components and antioxidative effects of Solanum lyratum Thunb. Kor. J. Pharmacogn. 26: (in Korean). Shin, H.C Technique for container seedlings of warmtemperate tree species. Proceedings of Korean 4th Technique Seminar of KCSRS. pp (in Korean). Shin, M.O., J.H. Park and J.O. Moon Effect of Solanum lyratum extract on dimethylniatrosamine-induced liver damage in rats. Kor. J. Pharmacogn. 34:60-64 (in Korean). Shin, M.O The effects on antimicrobial and cytotoxicity of Solanum lyratum fractions. Kor. J. Life Sci. 15: (in Korean). Spicer, P.B. and L.A. Dionne Use of gibberellin to hasten germination of Solanum seed. Nature 189: Sun, L.X., W.W. Fu, W. Li, K.S. Bi and M.W. Wang Diosgenin glucuronides from Solanum lyratum and their cytotoxicity against tumor cell lines. Zeitschrift fur Naturforschung Section C, Biosciences 61: Tzortzakis, N.G Effect of pre-sowing treatment on seed germination and seedling vigour in endive and chicory. Hort. Sci. (Prague). 36: Yang, J.S., C.C Wu, C.L. Kuo, Y.H. Lan, C.C. Yeh, C.C. Yu, J.C. Lien, Y.M. Hsu, W.W. Kuo, W.G. Wood, M. Tsuzuki, and J.G. Chung Solanum lyratum extracts induce extrinsic and intrinsic pathways of apoptosis in WEHI-3 murine leukemia cells and inhibit allograft tumor. Evid Based Complement Altern Med. 2012:ID Yeates, S.J., G.A. Constable and T.M Cumstie Developing management options for mepiquat chloride in tropical winter seasoncotton. Field Crops Res. 74: (Received 21 April 2014 ; Revised 21 May 2014 ; Accepted 28 May 2014) -370-

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386-390.hwp 386 HANYANG MEDICAL REVIEWS Vol. 29 No. 4, 2009 우리나라 미숙아의 통계와 의료비용 Statistics and Medical Cost of Preterm in Korea 윤혜선 을지대학교 노원을지병원 소아청소년과학교실 Hye Sun Yoon, M.D., Ph.D., Department of Pediatrics, Nowon