Leaf anatomy, gas exchange and photosynthetic enzyme activity in Flaveria kochiana
Erika A. Sudderth A D , Riyadh M. Muhaidat B , Athena D. McKown B , Ferit Kocacinar C and Rowan F. Sage BA Department of Organismic and Evolutionary Biology, Harvard University, Biological Laboratories, 16 Divinity Ave., Cambridge, MA 02138 11, USA.
B Department of Ecology and Evolutionary Biology, University of Toronto, Toronto, Ontario, M5S1A1, Canada.
C Faculty of Forestry, Kahramanmaras Sutcu Imam University, 46100 Kahramanmaras, Turkey.
D Corresponding author. Email: sudderth@fas.harvard.edu
Functional Plant Biology 34(2) 118-129 https://doi.org/10.1071/FP06263
Submitted: 19 October 2006 Accepted: 4 December 2006 Published: 12 February 2007
Abstract
Flaveria (Asteraceae) is one of the few genera known to contain both C3 and C4 species, in addition to numerous biochemically-intermediate species. C3-C4 and C4-like intermediate photosynthesis have arisen more than once in different phylogenetic clades of Flaveria. Here, we characterise for the first time the photosynthetic pathway of the recently described species Flaveria kochiana B.L. Turner. We examined leaf anatomy, activity and localisation of key photosynthetic enzymes, and gas exchange characteristics and compared these trait values with those from related C4 and C4-like Flaveria species. F. kochiana has Kranz anatomy that is typical of other C4 Flaveria species. As in the other C4 lineages within the Flaveria genus, the primary decarboxylating enzyme is NADP-malic enzyme. Immunolocalisation of the major C4 cycle enzymes, PEP carboxylase and pyruvate, orthophosphate dikinase, were restricted to the mesophyll, while Rubisco was largely localised to the bundle sheath. Gas exchange analysis demonstrated that F. kochiana operates a fully functional C4 pathway with little sensitivity to ambient oxygen levels. The CO2 compensation point (2.2 µbar) was typical for C4 species, and the O2-response of the CO2 compensation point was the same as the C4 species F. trinervia. Notably, F. vaginata (B.L. Robinson & Greenman), a putative C4-like species that is the nearest relative of F. kochiana, had an identical response of the CO2 compensation point to O2. Furthermore, F. vaginata, exhibited a carbon isotope ratio (–15.4‰) similar to C4 species including F. australasica Hooker, F. trinervia Spreng. C. Mohr and the newly characterised F. kochiana. F. vaginata could be considered a C4 species, but additional studies are necessary to confirm this hypothesis. In addition, our results show that F. kochiana uses an efficient C4 cycle, with the highest initial slope of the A/Ci curve of any C4 Flaveria species.
Additional keywords: C4 photosynthesis, C3-C4 photosynthesis, C4 evolution, CO2 compensation point, Kranz anatomy.
Acknowledgements
The authors thank Dr Espinosa-García for generous assistance coordinating our research activities, Dr Louis Villaseñor for providing locality information and specimen identification, and Martín Paredes Flores and Isabelle Blanckaert for assistance with the collection of F. kochiana specimens. We also thank the Secretaría de Medio Ambiente y Recursos Naturales and the Secretaría de Relaciones Exteriors of Mexico for granting permission to collect F. kochiana specimens. We appreciate the experimental advice and helpful discussions of the manuscript provided by Dr Nancy G. Dengler and Dr Noel M. Holbrook, and suggested revisions from two anonymous reviewers. This work was supported by NSF DDIG (DEB-0407716) and the Deland Fund (EAS) and NSERC (ADM and RM).
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