title: Sugarcane smut: shedding light on the development of the whip-shaped sorus.
abstract: Sugarcane smut is caused by the fungus Sporisorium scitamineum (Ustilaginales/Ustilaginomycotina/Basidiomycota), which is responsible for losses in sugarcane production worldwide. Infected plants show a profound metabolic modification resulting in the development of a whip-shaped structure (sorus) composed of a mixture of plant tissues and fungal hyphae. Within this structure, ustilospores develop and disseminate the disease. Despite the importance of this disease, a detailed histopathological analysis of the plant-pathogen interaction is lacking.
  
  
  
  The whip-shaped sorus was investigated using light microscopy, scanning and transmission electron microscopy, histochemical tests and epifluorescence microscopy coupled with deconvolution.
  
  
  
  Sorus growth is mediated by intercalary meristem activity at the base of the sorus, where the fungus causes partial host cell wall degradation and formation of intercellular spaces. Sporogenesis in S. scitamineum is thallic, with ustilospore initials in intercalary or terminal positions, and mostly restricted to the base of the sorus. Ustilospore maturation is centrifugal in relation to the ground parenchyma and occurs throughout the sorus median region. At the apex of the sorus, the fungus produces sterile cells and promotes host cell detachment. Hyphae are present throughout the central axis of the sorus (columella). The plant cell produces callose around the intracellular hyphae as well as inside the papillae at the infection site.
  
  
  
  The ontogeny of the whip-shaped sorus suggests that the fungus can cause the acropetal growth in the intercalary meristem. The sporogenesis of S. scitamineum was described in detail, demonstrating that the spores are formed exclusively at the base of the whip. Light was also shed on the nature of the sterile cells. The presence of the fungus alters the host cell wall composition, promotes its degradation and causes the release of some peripheral cells of the sorus. Finally, callose was observed around fungal hyphae in infected cells, suggesting that deposition of callose by the host may act as a structural response to fungal infection.
authors: Departamento de Genética, Escola Superior de Agricultura 'Luiz de Queiroz', Universidade de São Paulo, 13418-900 Piracicaba, Brazil.|Departamento de Ciências Biológicas, Escola Superior de Agricultura 'Luiz de Queiroz', Universidade de São Paulo, 13418-900 Piracicaba, Brazil.
authors: João Paulo R
authors: JPR
authors: Marques
authors: Departamento de Ciências Biológicas, Escola Superior de Agricultura 'Luiz de Queiroz', Universidade de São Paulo, 13418-900 Piracicaba, Brazil.
authors: Beatriz
authors: B
authors: Appezzato-da-Glória
authors: Department of Mycology, Cluster for Integrative Fungal Research (IPF), Goethe-Universität Frankfurt am Main, D-60438 Frankfurt am Main, Germany.
authors: Meike
authors: M
authors: Piepenbring
authors: Departamento de Fitopatologia e Nematologia, Escola Superior de Agricultura 'Luiz de Queiroz', Universidade de São Paulo, 13418-900 Piracicaba, Brazil.
authors: Nelson S
authors: NS
authors: Massola
authors: Departamento de Genética, Escola Superior de Agricultura 'Luiz de Queiroz', Universidade de São Paulo, 13418-900 Piracicaba, Brazil.
authors: Claudia B
authors: CB
authors: Monteiro-Vitorello
authors: Departamento de Genética, Escola Superior de Agricultura 'Luiz de Queiroz', Universidade de São Paulo, 13418-900 Piracicaba, Brazil.
authors: Maria Lucia Carneiro
authors: MLC
authors: Vieira
mesh_terms: D054884:Host-Pathogen Interactions; D025301:Hyphae / Q000502:physiology / Q000648:ultrastructure; D008855:Microscopy, Electron, Scanning; D046529:Microscopy, Electron, Transmission; D010935:Plant Diseases / Q000382:microbiology*; D031786:Saccharum / Q000382:microbiology*; D013172:Spores, Fungal / Q000254:growth & development / Q000502:physiology / Q000648:ultrastructure; D014587:Ustilaginales / Q000254:growth & development / Q000502:physiology* / Q000648:ultrastructure
keywords: Callose; Saccharum spp.; Sporisorium scitamineum; histopathological analysis; plant anatomy; smut disease