New Structural/Biochemical Associations in Leaf Blades of C4 Grasses (Poaceae)

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Abstract
Forty-three previously uninvestigated, mainly Australian, grass species (Poaceae) were assayed for activities of C4 acid decarboxylating enzymes (NADP-malic enzyme, NADP-ME; NAD-malic enzyme, NAD-ME; PEP carboxykinase, PCK). Twenty-five species exhibit long-established ('classical') associations between C4 type and structural features of leaf blade vascular bundles. However, Panicum virgatum and Triraphis mollis are NAD-ME species with structure like that of 'classical' PCK species. Seven Enneapogon species and Triodia scariosa are NAD-ME but are structurally intermediate between 'classical' NAD-ME and PCK species. Alloteropsis semialata (R.Br.) Hitch. is PCK, the first recorded non-NADP-ME XyMS - species, and Pheidochloa gracilis S.T. Blake and five Eriachne species are the first known XyMS+ NADP-ME species, with either centripetal or centrifugal/peripheral PCR (photosynthetic carbon reduction, or Kranz) cell chloroplasts. A suberised lamella is absent from the PCR cell walls of all species with an even PCR bundle sheath outline, irrespective of C4 type, as well as from NADP-ME Aristida, Eriachne and Pheidochloa; it is present in all other species with an uneven outline and centrifugal/peripheral chloroplasts. Pheidochloa gracilis and Eriachne spp. have unusually well-developed grana in PCR cell chloroplasts for NADP-ME species. This new-found structural/biochemical diversity is discussed in relation to high [CO2] maintenance in PCR cells.