title: Tapetum and middle layer control male fertility in Actinidia deliciosa.
abstract: OBJECTIVE
  Dioecism characterizes many crop species of economic value, including kiwifruit (Actinidia deliciosa). Kiwifruit male sterility occurs at the microspore stage. The cell walls of the microspores and the pollen of the male-sterile and male-fertile flowers, respectively, differ in glucose and galactose levels. In numerous plants, pollen formation involves normal functioning and degeneration timing of the tapetum, with calcium and carbohydrates provided by the tapetum essential for male fertility. The aim of this study was to determine whether the anther wall controls male fertility in kiwifruit, providing calcium and carbohydrates to the microspores.
  
  
  METHODS
  The events occurring in the anther wall and microspores of male-fertile and male-sterile anthers were investigated by analyses of light microscopy, epifluorescence, terminal deoxynucleotidyl transferase-mediated dUTP nick-end labelling (TUNEL assay) and transmission electron microscopy coupled with electron spectroscopy. The possibility that male sterility was related to anther tissue malfunctioning with regard to calcium/glucose/galactose provision to the microspores was also investigated by in vitro anther culture.
  
  
  RESULTS
  Both tapetum and the middle layer showed secretory activity and both degenerated by programmed cell death (PCD), but PCD was later in male-sterile than in male-fertile anthers. Calcium accumulated in cell walls of the middle layer and tapetum and in the exine of microspores and pollen, reaching higher levels in anther wall tissues and dead microspores of male-sterile anthers. A specific supply of glucose and calcium induced normal pollen formation in in vitro-cultured anthers of the male-sterile genotype.
  
  
  CONCLUSIONS
  The results show that male sterility in kiwifruit is induced by anther wall tissues through prolonged secretory activity caused by a delay in PCD, in the middle layer in particular. In vitro culture results support the sporophytic control of male fertility in kiwifruit and open the way to applications to overcome dioecism and optimize kiwifruit production.
authors: Department of Environmental Biology, 'Sapienza' University of Rome, Rome, Italy.
authors: Giuseppina
authors: G
authors: Falasca
authors: Simone
authors: S
authors: D'Angeli
authors: Rita
authors: R
authors: Biasi
authors: Laura
authors: L
authors: Fattorini
authors: Maja
authors: M
authors: Matteucci
authors: Antonella
authors: A
authors: Canini
authors: Maria Maddalena
authors: MM
authors: Altamura
mesh_terms: D029042:Actinidia / Q000166:cytology / Q000254:growth & development / Q000502:physiology*; D017209:Apoptosis / Q000502:physiology*; D002118:Calcium / Q000378:metabolism; D002473:Cell Wall / Q000378:metabolism; D035264:Flowers / Q000166:cytology / Q000254:growth & development / Q000502:physiology; D005947:Glucose / Q000378:metabolism; D046529:Microscopy, Electron, Transmission; D008856:Microscopy, Fluorescence; D051479:Plant Infertility / Q000502:physiology*; D011058:Pollen / Q000166:cytology / Q000254:growth & development / Q000502:physiology*; D012098:Reproduction
chemical_list: D005947:Glucose; D002118:Calcium
keywords: Actinidia deliciosa; anther; anther culture; calcium; dioecism; flower male fertility flower male sterility; glucose; middle layer; pollen; programmed cell death; tapetum
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