Developmental Anatomy and Branching of Roots of Four Zeylanidium Species (Podostemaceae), with Implications for Evolution of Foliose Roots
Y. HIYAMA1, I. TSUKAMOTO1, R. IMAICHI*,1 and M. KATO2
1 Department of Chemical and Biological Sciences, Faculty of Science, Japan Women’s University, 2-8-1 Mejirodai, Tokyo 112-8681, Japan and 2 Department of Biological Sciences, Graduate School of Science, University of Tokyo, 7-3-1 Hongo, Tokyo 113-0033, Japan
* For correspondence. Fax 03 5981 3658, e-mail [email protected]
Received: 25 June 2002; Returned for revision: 21 August 2002; Accepted: 16 September 2002 Published electronically: 24 October 2002
Podostemaceae have markedly specialized and diverse roots that are adapted to extreme habitats, such as seasonally submerged or exposed rocks in waterfalls and rapids. This paper describes the developmental anatomy of roots of four species of Zeylanidium, with emphasis on the unusual association between root branching and root-borne adventitious shoots. In Z. subulatum and Z. lichenoides with subcylindrical or ribbon-like roots, the apical meristem distal (exterior) to a shoot that is initiated within the meristem area reduces and loses meristematic activity. This results in a splitting into two meristems that separate the parental root and lateral root (anisotomous dichotomy). In Z. olivaceum with lobed foliose roots, shoots are initiated in the innermost zone of the marginal meristem, and similar, but delayed, meristem reduction usually occurs, producing a parenchyma exterior to shoots located between root lobes. In some extreme cases, due to meristem recovery, root lobing does not occur, so the margin is entire. In Z. maheshwarii with foliose roots, shoots are initiated proximal to the marginal meristem and there is no shoot–root lobe association. Results suggest that during evolution from subcylindrical or ribbon-like roots to foliose roots, reduction of meristem exterior to a shoot was delayed and then arrested as a result of inward shifting of the sites of shoot initiation. The evolutionary reappearance of a protective tissue or root cap in Z. olivaceum and Z. maheshwarii in the Zeylanidium clade is implied, taking into account the reported molecular phylogeny and root-cap development in Hydrobryum.
Key words: Branching, developmental anatomy, exogenous vs. endogenous origin of root, meristem, Podostemaceae, root, root cap, shoot, Zeylanidium.
Annals of Botany 90: 735-744, 2002. © 2002 Annals of Botany Company.