title: The unique pseudanthium of Actinodium (Myrtaceae) - morphological reinvestigation and possible regulation by CYCLOIDEA-like genes.
abstract: BACKGROUND
  Genes encoding TCP transcription factors, such as CYCLOIDEA-like (CYC-like) genes, are well known actors in the control of plant morphological development, particularly regarding the control of floral symmetry. Despite recent understanding that these genes play a role in establishing the architecture of inflorescences in the sunflower family (Asteraceae), where hundreds of finely organized flowers are arranged to mimic an individual flower, little is known about their function in the development of flower-like inflorescences across diverse phylogenetic groups. Here, we studied the head-like pseudanthium of the Australian swamp daisy Actinodium cunninghamii Schau. (Myrtaceae, the myrtle family), which consists of a cluster of fertile flowers surrounded by showy ray-shaped structures, to fully characterize its inflorescence development and to test whether CYC-like genes may participate in the control of its daisy-like flowering structures.
  
  
  RESULTS
  We used standard morphological and anatomical methods to analyze Actinodium inflorescence development. Furthermore, we isolated Actinodium CYC-like genes using degenerate PCR primers, and studied the expression patterns of these genes using quantitative RT-PCR. We found that the ray-shaped elements of Actinodium are not single flowers but instead branched short-shoots occasionally bearing flowers. We found differential expression of CYC-like genes across the pseudanthium of Actinodium, correlating with the showiness and branching pattern of the ray structures.
  
  
  CONCLUSIONS
  The Actinodium inflorescence represents a novel type of pseudanthium with proximal branches mimicking ray flowers. Expression patterns of CYC-like genes are suggestive of participation in the control of pseudanthium development, in a manner analogous to the distantly related Asteraceae. As such, flowering plants appear to have recruited CYC-like genes for heteromorphic inflorescence development at least twice during their evolutionary history.
authors: Institut für Spezielle Botanik und Botanischer Garten, Johannes Gutenberg-Universität, Mainz, 55099, Germany.
authors: Regine
authors: R
authors: Claßen-Bockhoff
authors: Department of Biological Sciences, University at Buffalo (SUNY), Buffalo, NY, USA.
authors: Raili
authors: R
authors: Ruonala
authors: Institut für Spezielle Botanik und Botanischer Garten, Johannes Gutenberg-Universität, Mainz, 55099, Germany.
authors: Kester
authors: K
authors: Bull-Hereñu
authors: Kings Park and Botanic Garden, West Perth, WA, 6005, Australia.
authors: Neville
authors: N
authors: Marchant
authors: Department of Biological Sciences, University at Buffalo (SUNY), Buffalo, NY, USA.
authors: Victor A
authors: VA
authors: Albert