식물병연구 Res. Plant Dis. 18(4) : 354 360 (2012) http://dx.doi.org/10.5423/rpd.2012.18.4.354 Research Article Open Access Research in Plant Disease The Korean Society of Plant Pathology 감귤녹색곰팡이병방제를위한천산용추출물의실용적적용 이지현 강성우 송정영 김홍기 * 충남대학교농업생명과학대학응용생물학과 Practical Application of Dioscorea quinqueloba Extract for the of Citrus Green Mold Ji Hyun Lee, Sung Woo Kang, Jeong Young Song and Hong Gi Kim* Department of Applied Biology, Chungnam National University, Daejeon 305-764, Korea (Received on May 14, 2012; Revised on November 14, 2012; Accepted on November 15, 2012) This study tested the antifungal compound obtained from a medicinal plant, Dioscorea quinqueloba Thunb., in order to search the possibility of practical application of this product in agriculture through evaluating its activity using the citrus fruits. The extract of D. quinqueloba Thunb., which has the strongest antifungal activity, was selected as a candidate among 101 plant extracts. Based on this examination concerning antifungal activity of the product on Penicillium digitatum in vitro, it was confirmed its effect of mycelial growth inhibition showed over 87% at 0.5 mg/ml concentration. This natural product showed the stability of the substance, as it was not significantly influenced by ph, temperature, or ultraviolet radiation. While citrus fruits were stored at room temperature, P. digitatum was inoculated into them in order to prepare a similar environmental conditions with epidemic occurrence of the mold. As the result of our investigation, the disease preventive effects of the active antifungal substance evidenced a 100% at 0.5 mg/ml. When the phytotoxicity of the selected natural product on citrus at 2 mg/ml was assessed, we noted no toxic effects. Based on the superior preventive effects from this natural product extracted from the plant, it is presumed to be very useful in agricultural applications for the control of green mold, P. digitatum, which has been occurred often the biggest problem in the storage of citrus fruits. Keywords : Biological control, Dioscorea quinqueloba Thunb., Penicillium digitatum, Plant extract, Storage disease 서론 Penicillium속균들에의한병들은작물의생육기간보다는수확후포장, 저장및운송되는과정중에상처를통해주로발생된다 (Paster와 Bullerman, 1988). 감귤류에서도녹색곰팡이병을일으키는 P. digitatum과푸른곰팡이병을일으키는 P. italicum에의한피해는심각하여저장기간동안발생하는손실의 90% 이상을차지하고있다 (Brown 과 Eckert, 2000). 특히 P. digitatum은 Penicillium속병원균들중에서가장강력한병원성을지니고있고, 전염력또한매우강해이병원균에의해경제적인손실이가장 *Corresponding author Phone) +82-42-821-5768, Fax) +82-42-823-8679 Email) hgkim@cnu.ac.kr 크게발생한다. 과거에는주요감귤저장병인녹색곰팡이병방제를위해유기합성농약인 imazalil, thiabendazole, pyrimethanil, fludioxonil과 sodium o-phenylphenate의단제및혼합제가효과적인방제제로널리사용되어졌다 (Larrigaudiére 등, 2002; Palou 등, 2001; Smilanick 등, 2006). 그러나 imazalil, thiabendazole과 sodium o-phenylphenate에대한약제저항성을나타내는 P. digitatum 균이발생함에따라효과적방제에문제점이제기되었다 (Fogliata 등, 2001; Kinay 등, 2007). 최근들어전세계적으로소비자의환경문제에대한인식이증가하고, 건강에대한관심또한매우높아지게되자소비자의요구를만족시키며환경에대한위험성을감소시킬수있는유기합성농약을대체할만한방제제개발이절실히요구되었다 (Holmes와 Eckert, 1999; Hao 등, 2011).
감귤녹색곰팡이병방제를위한천산용추출물의실용적적용 355 식물추출물과미생물을이용한생물학적방제는저독성이면서친환경적인방제법으로가장각광받으며다양한연구가시도되었으며 (Costa 등, 2001; Janisiewicz와 Korsten, 2002), 식물추출물로서는 essential oil이비교적많이분리되어광범위한항균활성효과가밝혀졌지만, 그대부분이식품첨가물로만이용될뿐작물의병방제를위해서는최근 methyl eugenol과 fennel, thyme 등이식물병원균에광범위한항균활성을보유한것으로보고되고있다 (Kim 등, 2009; Tzortzakis, 2007; Yahyazadeh, 2008). 또한과일의병에대한저항성을향상시키기위해식물생장을조절하는 jasmonic acid(ja; Droby 등, 1999), β- aminobutyric acid(baba; Porat 등, 1999b) 와같은천연화합물을이용하거나 ultraviolet(uv) (Droby 등, 1993; Porat 등, 1999a) 나뜨거운물세척 (Porat 등, 2000) 등의물리적인방제법에관한연구가수행된바있다. 최근에는그유용한특성들이밝혀지면서국내 외적으로식물유래천연물을이용한식물병방제가시도되고있으며그중요성과가능성또한점차부각되고있다. 그러나다양한식물체의종류에비해천연물의이용가치에관한연구는극히미미한실정임에도불구하고이또한식물병이나가축질병에대한방제제로서의연구보다는사람에대한약리작용의연구가중시되었던점을고려하면식물유래천연물질을이용한식물병방제방법의개발가능성은더욱높으며그의미가크다 (Mekbib 등, 2007). 특히세계적인추세에따라병해종합관리방법이인체에무해하고환경친화적인생물적방제법과함께이용되기때문에인체에무해한식물유래친환경천연물농약의개발은필수적인상황이다. 단풍마 (Dioscorea quinqueloba Thunb.) 의뿌리인천산용은다수의 steroid saponin을함유하고있으며 (Sautour 등, 2007), 그동안한약제나건강식품으로널리사용되어져왔다 (Kum 등, 2006). 그러나아직까지그로부터항균물질이추출되어식물병해방제제등농업용으로사용된적은없다. 이연구는국내감귤저장에서가장큰피해를주고있는 P. digitatum에의한감귤녹색곰팡이병방제를위하여 P. digitatum에항균활성을보이는약용식물천산용 (Dioscorea quinqueloba Thunb.) 추출물의현장적용가능성을확인하고자실시하였다. 재료및방법 최적식물체추출물선발. 충남대학교식물병리학연구실에추출, 보관중이거나국립원예특작과학원인삼특 작부인삼약초가공팀으로부터분양받은총 101가지의 methanol 식물추출물을본실험에시료로사용하였다 (Table 1). 식물추출물의항균활성을검정하기위해공시균주 P. digitatum KACC 41898을 potato dextrose agar(pda, Difco Co.) 배지에접종하여 25 o C에서 7일간배양한다음멸균수에포자를수거하여포자현탁액을준비하였다. 이를이용하여 PDA 100 ml당 6 10 5 개의포자를함유하는혼합배지를만든후식물추출물을점적하여건조시킨 paper disc(8 mm, ADVANTEC ) 를치상하고 25 o C에서 7일간배양하여병원균에대한저지원의지름을측정, 가장강한활성을보인천산용 (Dioscorea quinqueloba Thunb.) 추출물을선발하였다. 천산용추출물의 In vitro 상 P. digitatum 억제효과. 감귤저장병균 P. digitatum에대한천산용추출물의농도별방제효과를확인하기위해각각 0.05, 0.1, 0.2, 0.3, 0.4, 0.5 mg/ml 농도의천산용추출물원액을함유한 PDA 배지를만들고대조구는다른성분이함유되지않은 PDA 배지를사용하였다. 배지중앙에 P. digitatum을접종한다음 25 o C에서 2주간배양시킨균사의균사생장정도를측정하고생장억제율을조사하였다. 천산용추출물의안정성및특성조사. 항균물질의 ph 에대한안정성을알아보기위하여 ph 3.0 7.0로조절된 PDA 배지에 0.2 mg/ml의천산용추출물을혼합하여분주한후 P. digitatum KACC 41898을접종하여균사생육억제정도를검정하였으며, 대조구는 ph를조절하지않은 PDA 배지를사용하였다. 항균활성물질의온도에대한안정성을밝히고자천산용추출물 0.2 mg/ml을함유한 PDA 배지를 121 o C, 20분간고압멸균한후 P. digitatum을접종하여균사생육억제활성을분석하였는데배지온도가 50 o C일때동일한농도의활성물질을혼합한대조구와비교하여항균특성의변화여부를분석하였다. UV에대한안정성을확인하기위해 UV와햇빛에서 24시간노출시킨활성물질 0.2 mg/ml을 PDA 배지에혼합하여 P. digitatum을접종한다음, 균사생육억제활성을검정하였다. UV 및햇빛에노출되지않은활성물질을같은농도로혼합한 PDA배지를대조구로이용하였다. 천산용추출물을점적하여건조시킨 paper disc를 petri dish 뚜껑안쪽에부착시켜 P. digitatum에대한균사생육억제율을조사하여휘발성유무를확인하였다. 저장병자연발생시물질농도별방제효과. 저장병의자연발생상태하에서천산용추출물의농도별방제효과를확인하고자활성물질을각각 0.1, 0.2, 0.3, 0.4, 0.5 mg/ ml 농도로 EtOH 10% 에희석하여준비하였다. 상처가없
356 이지현 강성우 송정영 김홍기 Table 1. Antifungal activity of plant extracts against Penicillium digitatum Scientific name Inhibition Inhibition zone a Scientific name zone a Datura stramonium var. chalybea Koch Senna occidentalis L. Lysimachia barystachys Bunge Angelica acutiloba Kitagawa Amorpha fruticosa L. Lespedeza cuneata G. Don Acer tataricum subsp. ginnala (Maxim.) Wesm. Albizia julibrissin Durazz. + Brassica campestris subsp. napus var. nippo-oleifera Asparagus officinalis L. Makino Artemisia dracunculus Potentilla chinensis Ser. Artemisia capillaris Thunb. Patrinia rupestris (Pall.) Juss. Lespedeza bicolor Turcz. Hibiscus manihot L. Artemisia japonica var. manshurica Kitam. Aruncus dioicus var. kamtschaticus Hara Artemisia sieversiana Ehrh. ex Willd. Glycyrrhiza glabra L. Lonicera maackii (Rupr.) Maxim. Dianthus japonicus Thunb. Vicia unijuga A. Braun Boehmeria tricuspis (Hance) Makino Thalictrum aquilegifolium var. sibiricum Solanum nigrum L. var. nigrum ++ Hieracium umbellatum L. Rubia cordifolia var. pratensis Maxim. Acer triflorum Kom. Rubus coreanus Miq. Rubia akane Nakai Persicaria thunbergii (Siebold & Zucc.) Potentilla supina L. Psoralea corylifolia L. Sorbaria sorbifolia var. stellipila Maxim. Euphorbia pekinensis Rupr. Alnus sibirica Fisch. ex Turcz. Scabiosa tschiliensis Cornus controversa Hemsl. ex Prain Penthorum chinense Pursh + Eleutherococcus divaricatus var. chiisanensis (Nakai) Draba nemorosa L. Cleyera japonica Thunb. Verbesina alternifolia Britton Quercus myrsinaefolia Blume Serratula coronata var. Insularis Kitamura Suaeda maritima (L.) Dumort. Rudbeckia laciniata L. Betula platyphylla var. japonica (Miq.) Hara Aster tartaricus L. Ambrosia trifida L. var. trifida Achillea alpina var. discoidea (Regel) Kitam. Linum usitatissimum L. Ipomea aquatica Forsk Senecio vulgaris L. Campsis grandiflora (Thunb.) K. Loisel Cudrania tricuspidata (Carr.) Zelkova serrata (Thunb.) Makino Saururus chinensis (Lour.) Baill. Asparagus oligoclonos Maxim. Silene armeria L. Viola patrinii DC. Abelomoschus esculentus Valeriana officinalis var. latifolia MIQ. Chrysanthemum burbankii Makino Patrinia saniculaefolia Hemsl. ++ Tanacetum bungeia Angelica gigas Nakai Stevia rebaudiana Bertoni Coriandrum astivum L. Platycodon grandiflorum A.D.C. Angelica dahurica Momordica charantia L. Oenanthe javanica (Blume) DC. Althaea rosea Cav. Bupleurum falcatum L. Synurus deltoides (Aiton) Nakai Physalis alkekengi L. var. franchetii Mort Achyranthes japonica (Miq.) Nakai Rehmannia glutinosa (Gaertner) Liboschitz Gomphrena globosa L. Astilbe chinensis var. coreana Rosa multiflora Thunb. var. multiflora Phellodendron amurense Rupr. Polygonatum odoratum var. pluriflorum Ruta graveolens L. Taraxacum officinale Weber Geum japonicum Thunb. Campanula takesimana Nakai Clematis apiifolia DC. Equisetum arvense Linne Lysimachia clethroides Duby Prunus mume Sieb. et Zucc Lysimachia vulgaris var. davurica (Led.) R. Knuth. Sedum sarmentosum Bunge Hibiscus trionum L. Vitex rotundifolia L. fil. Geranium krameri Fr. et Sav. Cornus officinalis Sieb et Zucc Dioscorea quinqueloba Thunb. +++ Caryopteris incana (Thunb.) Miq. a Growth inhibition was determined on 7 days after incubation at 25 o C. Inhibition zone; : 0 mm, +: <10 mm, ++: <20 mm, +++: 20 mm. > =
감귤녹색곰팡이병방제를위한천산용추출물의실용적적용 357 는감귤 ( 품종 : 온주밀감 ) 에농도별물질제제 stock을감귤 1개당 1ml씩분무처리하였고, 처리구당 15반복씩수행하였다. 대조약제는현재시판되고있는 iminoctadine tris와 prochloraz manganese complex를각각 1,000배농도로희석하여사용하였다. 감귤은밀봉한상태로온도와습도가유지되는발병최적조건의실온 (22 o C) 에서 10일간자연발생시감귤의녹색곰팡이병에대한방제효과를조사하였으며 2차례에걸쳐 3반복수행하였다. 병원균접종후감귤저장병방제효과. 병원균이접종된상태에서의방제효과를확인하고자 P. digitatum을 PDA 배지에접종하여 25 o C에서 2주간키운후멸균수로포자를수거하여 1 10 3, 1 10 4, 1 10 5 spores/ml로조정하였으며접종원을감귤에분무접종한다음 22 o C와포화습도를유지시켜발생최적의환경을만들어주었다. 균접종 7시간경과후, 0.3, 0.4, 0.5 mg/ml 농도의활성물질제제 stock을감귤에분무살포하였다. 표면의물기를자연건조시킨다음시판중인감귤상자에넣어실험이진행되는동안습도를충분히유지시킨상태에서 22 o C로보관하였다. 활성물질제제는 1회살포하였으며, 실험은 3반복으로수행하였다. 약해검정. 과실에대한약해발생유무를검정하기위하여 0.5% 차아염소산나트륨 (NaOCl) 에 3분간표면소독한감귤에 1mg/ml과배량인 2mg/ml 농도의활성물질을감귤과실에분무처리하였다. 3, 5, 7일후에달관조사방법으로약해유무를조사하였고, 각처리는 3반복으로수행되었다. 결과및고찰 항균성식물추출물선발. 총 101가지식물체로부터얻은 methanol 추출물의감귤녹색곰팡이병균 P. digitatum KACC 41898에대한항균활성을검정한결과, 천산용추출물이 20 mm 이상의큰저지원을형성하여아주강력한항균활성을보였으며, 자귀나무, 낙지다리추출물은 10 20 mm 억제하였고, 까마중, 금마타리는 10 mm 미만의약한항균활성이확인되었다 (Table 1). 따라서 P. digitatum에가장강력한항균활성을나타낸천산용을본실험의재료로선발하였다. 천산용은마속에속하는단풍마 (D. quinqueloba Thunb.) 의뿌리로서마속의식물은 50종류이상의 steroid saponin을함유하고있으며 (Sautour 등, 2007), 오랜동안민간요법치료제나건강식품으로는널리사용되어져왔으나 (Kum 등, 2006), 아직까지감귤녹색곰팡이병을비롯하여각종식물병해방제를위해농업용으로사용된적은없었다. Fig. 1. Inhibition rates of mycelial growth of Penicillium digitatum on PDA media containing different concentration of an antifungal compound extracted from Dioscorea quinqueloba. 천산용추출물의 In vitro 상 P. digitatum 억제효과. 항균활성물질이각각 0.05, 0.1, 0.2, 0.3, 0.4, 0.5 mg/ml의낮은농도로첨가된배지를만들어 P. digitatum 의균사생장억제율을측정한경우, 0.1 mg/ml에서 50% 이상, 0.3 mg/ ml에서 70% 정도의균사생장억제율이확인되었고, 0.5 mg/ml에서는 85% 이상강한균사생장이억제되었다 (Fig. 1). 식물추출물을이용한감귤저장병방제에대한연구보고로는 Tzortzakis와 Economakis(2007) 에의해 lemongrass essential oil 0.5 mg/ml 이상의농도에서 P. digitatum의포자발아를완벽하게억제하였다. 이로써 P. digitatum 방제물질로서뛰어난효과를보인천산용추출물은감귤녹색곰팡이병균에대해효과적인방제제로서충분한가능성을가지고있음을확인할수있었다. 항균활성물질의안정성및휘발성조사. 선발된천산용추출물의 ph에대한안정성을조사한결과 ph 3 7의넓은범위에서 P. digitatum에항균활성을유지하였기때문에 ph에안정함이확인되었다. 또한항균활성물질을 121 o C에서 20분간처리하여열에의한항균활성의변화를조사했을때에도활성이유지되었으므로이물질은열에매우안정적임을알수있었으며, UV와햇빛처리에도항균활성의변화가전혀일어나지않았다 (Fig. 2). 그리고휘발성확인실험에서는항균활성을보이지않았으므로비휘발성물질로판단되었다. 이물질은 ph, 열처 리시에도활성의변화가없었으므로 ( 자료미제시 ) 매우안정한구조의화합물임을알수있었으나추후보다정확한물질규명이요망된다. ph, 온도, 광의영향에도안정적으로활성이유지되는본항균활성물질의특성은추후천연물농약으로서의개발및제품생산에매우유용하게작용하여제형화과정에서변함없이강한항균활성유지가가능할것이며, 제품화된이후보관과정에서도활성을잃지않고그대로유지할수있어상당히긴유효기간을가질것으로판단된다.
358 이지현 강성우 송정영 김홍기 Table 3. Disease control efficacy of antifungal compound extracted from Dioscorea quinqueloba against Penicillium digitatum treated with different concentration of the pathogen on citrus stored at 22 o C for 10 days Fig. 2. Stability concerning antifungal activity on Penicillium digitatum of methanol extract of Dioscorea quinqueloba against UV or light treatment. Table 2. effects against Penicillium digitatum by using different concentration of antifungal compound extracted from Dioscorea quinqueloba on citrus fruits Treatment Antifungal compound Diseased Concentration rate (mg/ml) value 0.05 33.3b a 20.1 0.1 33.3b 20.1 0.2 8.3a 80.1 0.3 0a 100 Iminoctadine tris 1 0a 100 Prochloraz manganese 1 6.7a 83.9 complex None treated 41.7b a Means followed by the same letter within the column are not significantly different by Duncan s multiple range test at 5% level. 자연발생시감귤저장병의농도별방제효과. 시판중인감귤 ( 품종 : 온주밀감 ) 에자연적으로발생하는녹색곰팡이병에대한활성물질의방제효과를검정하였을때, 0.2 mg/ml 처리구에서 80.1% 의높은방제가를보였고, 0.3 mg/ml에서는 100% 완벽하게방제되었다 (Table 2). 자연발생을유도한후물질을처리하였을때 0.2 mg/ml의저농도에서 80% 이상의높은방제가를얻은것으로보아항균물질은추후방제약제로활용시저농도처리만으로도충분한방제효과를볼수있으며, 제품화할경우에도소량의원자재만으로사용이가능함에따라큰어려움없이추후농업적인활용이가능할것으로판단된다. 본연구의활성물질은현재시판중인농약 iminoctadine tris 수화제 ( 벨쿠트, 경농 ), prochloraz manganese complex 수화제 ( 스포르곤, 한국삼공 ) 와유사한강한방제효과를보였으므로추후친환경적인천연물농약으로서의활용가 Number of P. digitatum treated (spores/ml) Extract concentration (mg/ml) Diseased rate value 1 10 3 0.4 0 100 0.3 4.4 75.3 0.5 0 100 0.3 17.8 52.9 1 10 4 0.4 11.1 70.6 0.5 0 100 0.3 24.4 47.8 1 10 5 0.4 13.3 71.5 0.5 2.2 95.3 1 10 3 17.8 1 10 4 37.8 1 10 5 46.7 능성이다시한번확인되었다. 병원균접종후감귤저장병방제효과. 병원균을인공접종하고병발생에호조건을만들어준환경에서활성물질의병방제효과검정을수행한결과 5일째에는모든처리구에서병의발생이미미하였고 0.5 mg/ml 농도처리구는전혀병이발생하지못하였다. 10일경과후방제효과는병원균 1 10 4 spores/ml 농도로접종한처리구는 0.4 mg/ml에서 70% 정도의방제효과를보였으며, 0.5 mg/ ml에서는방제가 100% 로완벽하게방제되었다. 1 10 5 spores/ml의고농도로접종시에는무처리구에서 46.7% 발병한것에비해, 0.5 mg/ml에서는단지 2.2% 만이발병함으로서방제가가 95.3% 나되었다 (Table 3). 이처럼자연상태에비해고농도로 P. digitatum 균을접종하여병발생에매우유리하도록환경을조성하였음에도불구하고, 0.5 mg/ml 처리에의해아주효과적으로방제가가능하였다. P. expansum에의한사과푸른곰팡이병을억제하는생강추출물, P. italicum의방제가가능한대극추출물및 essential oil을이용한 P. digitatum의방제등다양한항균성식물추출물을이용하여 Penicillum 속에의한저장병방제가보고된바있으나 in vitro 상에서의효과만검정되거나실제적인작물의저장시 Penicillum속병원균에대한억제효과는미미하였다 (Al-Mughrabi, 2003; Paik 등, 1997; Yahyazadeh, 2008). 본연구에서사용된천산용추출물은현실적으로감귤
감귤녹색곰팡이병방제를위한천산용추출물의실용적적용 359 처리하였을때 0.3 mg/ml에서완벽하게병이방제되었으며인위적으로고농도의 P. digitatum을접종한경우에도 0.5 mg/ml 농도에서 100% 의방제효과를보였다. 이천연항균물질을 2mg/ml 농도로감귤과실에처리하여도약해가나타나지않았으므로, 약해가없고방제효과가우수한이식물유래천연물질은감귤저장중에발생하는녹색곰팡이병방제를위해추후매우효과적인농업적활용이가능할것으로판단된다. Acknowledgments Fig. 3. Phytotoxicity test of the antifungal compound extracted from Dioscorea quinqueloba against citrus fruits. (A) 1 mg/ml treatment. (B) 2 mg/ml treatment. 의유통과정중최적의녹색곰팡이병발생조건이형성된다할지라도활성물질 0.5 mg/ml 농도에서매우효과적인병방제가가능하여그경제적손실을크게줄일수있을것으로보이는바추후친환경적방제제로서의활용가능성이충분하다고판단된다. 약해조사. 활성물질의과실에대한약해유무검정시 1mg/ml과 2mg/ml의고농도로처리했음에도 7일이지난후에도과실에무름, 반점, 탈색및당질과같은성분의용탈등이일어나지않아외관상약해가전혀없음을확인하였다 (Fig. 3). 따라서생물농약으로사용할때실수로인해고농도로살포될지라도물질처리에의해과실에피해가전혀일어나지않을것으로생각된다. 요약 총 101종의식물추출물가운데약용식물인천산용 (Dioscorea quinqueloba Thunb.) 추출물이가장강한항균활성을보유한것으로나타나감귤녹색곰팡이병 (Penicillium digitatum) 방제를위한실질적인농업적현장적용가능성을탐색하였다. 선발된천산용추출물은 PDA 배지에 0.5 mg/ml을처리하였을때, P. digitatum의균사생장을 87% 억제하였다. 그리고추출된항균활성물질은 ph, 온도, UV에영향을받지않는매우안정한물질임이확인되었다. 감귤저장상태하에서녹색곰팡이병방제를검정한결과, 자연적인병발생을유도한후항균활성물질을 This study received a number of plant extracts from Ginseng Research Division, National Institute of Horticultural Herbal Science and was conducted as a part of R&D projects of Korea Institute of Planning and Evalution for Technology of Food, Agriculture, Forestry and Fisheries (IPET) (Project No. 109065-3). References Al-Mughrabi, K. 2003. Antimicrobial activity of extracts from leaves, stems and flowers of Euphorbia macroclada against plant pathogenic fungi. Phytopathol. Mediterr. 42: 245 250. Brown, G. E. and Eckert, J. W. 2000. Penicillium Decays. In compendium of Citrus Diseases 2nd. Timmer, L. W., Garnsey, S. M. and Graham, J. H., eds. The American Phytopathological Society, St. Paul, Mn. 41 42. Costa, E., Teixidó, N., Usall, J., Atarés, E. and Viñas, I. 2001. Production of the biocontrol agent Pantoea agglomerans strain CPA-2 using commercial products and by-products. Appl. Microbiol. Biotechnol. 56: 367 371. Droby, S., Chalutz, E., Horev, B., Cohen, L., Gaba, V., Wilson, C. and Wisniewski, M. 1993. Factors affecting UV-induced resistance in grapefruit against the green mould decay caused by Penicillium digitatum. Plant Pathol. 42: 418 424. Droby, S., Porat, R., Cohen, L., Weiss, B., Shapira, B., Philosoph- Hadas, S. and Meir, S. 1999. Suppressing green mold decay in grape fruit with postharvest jasmonates application. J. Am. Soc. Hort. Sci. 124: 184 188. Fogliata, G. M., Torres, L. G. J. and Ploper, L. D. 2001. Detection of imazalil-resistant strains of Penicillium digitatum Sacc. In citrus packing houses of Tacuman Province (Argentina) and their behaviour against current employed and alternative fungicides. Rev. Ind. Agric. Tacuman. 77: 71 75. Hao, W., Li, H., Hu, M., Yang, L. and Rizwan-ul-Haq, M. 2011. Integrated control of citrus green and blue mold and sour rot by Bacillus amyloliquefaciens in combination with tea saponin. Postharvest Biol. Technol. 59: 316 323.
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