title: B- and C-class gene expression during corona development of the blue passionflower (Passiflora caerulea, Passifloraceae).
abstract: OBJECTIVE
  The origin of the passionflower corona, a complex series of structures between the petals and stamens, has intrigued botanists for centuries, but has proven intractable using traditional approaches. Supplementing developmental data with expression analyses of three floral identity genes, we test whether the corona in Passiflora caerulea (blue passionflower) is homologous to petals or stamens or whether an alternative hypothesis of the corona as a novel structure is supported.
  
  
  METHODS
  Corona development was investigated using scanning electron microscopy. Expression of the P. caerulea B-class genes PISTILLATA (PcPI) and TOMATO MADS6 (PcTM6), and C-class gene AGAMOUS (PcAG) was investigated using a combination of RT-PCR and mRNA in situ hybridization analyses.
  
  
  RESULTS
  Corona development starts as a ring of tissue at the base of petals. The outer radii and operculum initiate first at the periphery, followed by the inner radii and pali toward the center, and finally an annulus beneath the operculum. Late in development, a limen, the innermost component of the corona, develops from the side of the androgynophore. RT-PCR analyses indicate that the B-class genes PcPI and PcTM6 and C-class gene PcAG were all expressed in mature coronas. However, mRNA in situ hybridization analyses revealed complex temporal patterns of gene expression in the different corona elements.
  
  
  CONCLUSIONS
  Our data support the hypothesis that the corona is a composite structure, with the radii, pali, and operculum homologous to stamens, and the limen, which only expresses PcTM6, considered to be a novel structure distinct from the androgynophore.
authors: Department of Biological Sciences, California State University-Long Beach, 1250 Bellflower Boulevard, Long Beach, California 90840, USA.
authors: Claire A
authors: CA
authors: Hemingway
authors: Ashley R
authors: AR
authors: Christensen
authors: Simon T
authors: ST
authors: Malcomber