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Alternaria Black Spot of Crucifers: Symptoms, Importance of Disease, and Perspectives of Resistance Breeding

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Alternaria Black Spot of Crucifers: Symptoms, Importance of Disease, and Perspectives of Resistance Breeding 2012 vol. 76, 5-19 DOI: 10.2478/v10032-012-0001-6 ________________________________________________________________________________________ ALTERNARIA BLACK SPOT OF CRUCIFERS: SYMPTOMS, IMPORTANCE OF DISEASE, AND PERSPECTIVES OF RESISTANCE BREEDING Marcin NOWICKI, Marzena NOWAKOWSKA, Anna NIEZGODA, Elżbieta U. KOZIK Research Institute of Horticulture Konstytucji 3 Maja 1/3, 96-100 Skierniewice, Poland Received: June 6, 2012; Accepted: July 16, 2012 Summary Alternaria black spot of cruciferous vegetables, incited by different species of Alternaria, remains an increasing threat to Brassicaceae crops throughout the world, including Poland. Brassica plants are attacked by conidia of A. brassicae (Berk.) Sacc., A. brassicicola (Schw.) Wiltsh., A. raphani Groves & Skolko, and A. alternata (Fr.) Kreissler. The pathogens have a wide spectrum of hosts, such as head cabbage, Chinese cabbage, cauliflower, broccoli, and other crucifers includ- ing cultivated and wild grown plants. Alternaria pathogens usually cause damp- ing-off of seedlings, spotting of leaves of cabbages, blackleg of heads of cabbag- es, and spotting of cauliflower curds and broccoli florets. In oilseed rape, A. brassicae is the dominant invasive species, while in the cruciferous vegetables, both species, A. brassicae, and A. brassicicola are encountered. Infected seeds with spores on the seed coat or mycelium under the seed coat are the main means of distribution for these pathogens. The fungus can overwinter on susceptible weeds or crop debris and on seed plants, as well as on stecklings. Methods for disease prevention and control are based on combining agri- cultural management practices with chemical control. Using disease-free seeds or seeds treated with fungicides can greatly reduce disease incidence. After appearance of the first symptoms of disease, stringent fungicide spray program is an effective way to reduce losses. Many authors seem to agree, that the most economically feasible method of disease control is the development of resistant Brassicaceae crops varieties, as transgenic approach proved unsuccessful. Due to our increasing understanding of pathogen-host plant interactions, identifica- tion of resistance sources, and assessment of the resistance trait inheritance mode, breeding programs of Brassica crops for Alternaria resistance can be enhanced. This is of particular importance since recent years experience dynam- ic development of ecological and integrated plant production with an emphasis on plant biotic stress resistance. Highly resistant genetic resources have not been reported in Brassica cultivated species, although some varieties differ in their resistance/susceptibility level. Corresponding author: e-mail: [email protected] © Copyright by InHort 6 VEGETABLE CROPS RESEARCH BULLETIN 76 _____________________________________________________________________________________________________ Strong cross-incompatibility, polygenic background of the resistance (ad- ditive and dominant gene interactions), as well as the differences in ploidy be- tween the Brassica species of interest, render the transfer of Alternaria re- sistance from the wild species into the cultivated forms difficult. Additionally, it is often connected with employment of in vitro hybridization techniques, in- cluding somatic hybridization, embryo and ovary rescue, or protoplast fusion. key words: Alternaria spp., cruciferous plants, resistance, testing methods INTRODUCTION A. brassicicola and A. brassicae cause severe economic losses in sev- Cruciferous plants (Brassicaceae) eral different ways (Humpherson- worldwide are severely affected by the Jones & Maude 1982, Humpherson- Alternaria fungi. A. brassicae (Berk.) Jones 1989). Seed infection causes Sacc., A. brassicicola (Schw.) Wiltsh., reduced germination and seedling A. raphani Groves and Skolko, and A. vigour, in addition to pre- and post- alternata (Fr.) Kreissler belong to spe- emergence damping-off, and affects cies of major negative influence on the sale and use of infected/infested Brassicaceae plants. Although both, seed. Lesions on leaves, stems, and A. brassicicola and A. brassicae occur siliques reduce the photosynthetic on oleiferous and vegetable (olera- area and accelerate senescence in the ceous) Brassicas, the former is the dom- plant. These pathogens are responsi- inant invasive species of the vegetable ble for major seed yield losses in the Brassicas, while the oleiferous cruci- oleraceous Brassicas and this is the fers are primary hosts for the latter most important component of their fungus (Maude & Humpherson-Jones economic impact. The unsightly cos- 1980, Humpherson-Jones 1989). metic blemishing or rotting of the Both most common Alternaria head or wrapper leaves in vegetable pathogens usually cause black spot Brassicas as a consequence of patho- disease, manifested by damping-off of gens’ toxins and disease causes seedlings, spotting of leaves of cab- downgrading and crop losses in both bages, blackleg of heads of cabbages fresh and stored produce. A. bras- (head cabbage and Chinese), and spot- sicicola often occurs in conjunction ting/browning of cauliflower curds with A. brassicae and some other and broccoli florets. Black spot is the pathogens of the Brassicaceae. This most common disease in the crucifers confounds precise estimates of losses plantations located in tropical and caused individually by this pathogen sub-tropical regions. Due to weather in the field. conditions conducive to infection (see In Europe, alone the seed losses below), however, the disease consti- due to both pathogens were estimated tutes a serious problem in crucifers at up to 86% in B. oleracea in several plant production in long-lasting high years (Maude & Hampherson-Jones humidity or intensive rains regions, 1980, Humpherson-Jones 1989). Un- including Poland. der Polish climate conditions, the disease takes particularly high toll on M. NOWICKI et al. – ALTERNARIA BLACK SPOT … 7 __________________________________________________________________________________________________ the late and medium-late varieties of active ingredients (Amistar 250 SC, head-cabbage grown for sauerkraut Signum 33 WG, Zato 50 WG) and processing, as well as for storage. fungicides based on iprodione (Rovral Cabbage heads with characteristic FLO 255 SC) (Maude et al. 1984, symptoms of Alternaria black spot do Survilienė et al. 2010). This method, not store well, and their processing however, carries an economic disad- value is very low. This translates onto vantage and may prove ineffective significant economic impact of the under pathogen infection-conducive disease: In 2010, Polish cabbages and weather conditions, particularly other Brassicas production ranked 7th, among the seed crops. An alternative while cauliflower and broccoli ranked protection method to be employed is 8th globally (1141200 tonnes and use of antagonistic fungi; deployment 252325 tonnes, respectively) with net of Aureobasidium pullulans and Epi- worth of production of Int.$171 M and coccum nigrum on the crucifers leaves Int.$60,5 M, respectively (FAOSTAT: reduced the infection level under con- http://faostat.fao.org). trolled conditions (Pace & Campbell Alternaria prevention and con- 1974). Field studies concerning the trol methods include combining the biological control efficacy are yet to proper agro-technique with chemical be carried out. protection. An essential disease pre- Pathogen profile and infection pro- vention method is production of gress healthy seeds, obtained from planta- The current classification of Al- tions with heavy fungicide protection. ternaria fungi is as follows: Kingdom In the 2-year lasting crucifers seed Fungi, Phylum Ascomycota; Subdivi- production periods, good effects of sion Pezizomycotina; Class Dothide- protection against Alternaria infec- omycetes; Order Pleosporales; Family tions during the 1st year of growth Pleosporaceae; Subfamily mi-tosporic were expedited by fungicides contain- Pleosporaceae; Genus Al-ternaria ing iprodione as an active ingredient (http://www.uniprot.org/taxon- (Maude et al. 1984, Survilienė et al. omy/5598). There are 299 species listed 2010). In Poland, the only product in the genus (Kirk et al. 2008); most Al- containing it is Seed Protector T 75DS ternaria species are saprophytes that are WS (Zaprawa Nasienna). Since both commonly found in soil or on decaying pathogens survive on crop debris, plant tissues (Bart & Thomma 2003). seeds, and in association with weed Alternaria fungi proliferation is vege- hosts (Humpherson-Jones & Maude tative in character, and takes place by 1982, Humpherson-Jones 1989), crop means of conidial spores, airborne and debris management (for example found in the soil and water, as well as through crop rotation and deep tillage) indoors and on objects. Sexual re- and use of clean seed and proper weed combination (teleomorphy) occurs control should alleviate the disease. very rarely. After appearance of disease symp- Alternaria spp. may be grown on toms, one may achieve limitation of artificial agar media, with PDA (pota- the infection by repeated spray with to-dextrose agar) and V8 (V8 juive - fungicides containing strobilurines as agar) being the most popular in use. 8 VEGETABLE CROPS RESEARCH BULLETIN 76 _____________________________________________________________________________________________________ As observed in the in vitro cultures, Chinese cabbage, cauliflower, brocco- the pathogens develop fast-growing li, and of other crucifers. Infected thick colonies which are usually
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