Bienertia cycloptera Bunge ex Boiss., Chenopodiaceae, another C₄ Plant without Kranz Tissues[1]

Abstract

Following up an earlier study on Borszczowia aralocaspica, a second C₄ leaf type without Kranz tissues is described from Bienertia cycloptera Bunge ex Boiss. The species belongs also to tribe Suaedeae and grows in similar temporarily wet saline habitats in southwestern C Asia. Like Borszczowia, the species has the ability to perform C₄ photosynthesis in single chlorenchyma cells but the cytological compartmentation differs. Evidence is cited from anatomy (cytoplasmic compartmentation, starch formation), from carbon isotope composition and from indirect sources. It is documented by micrographs and δ¹³C values (mean - 14.7 ‰). Variations of the δ¹³C values subsequently found in newly formed leaves of the same greenhouse plants (- 15.5 to - 21.1 ‰) support the hypothesis that Bienertia is a facultative C₄/C₃ species. The new described bienertioid leaf type differs from the leaves of C₃ species in Suaeda by strict separation of the mesophyll into a central aqueous tissue without chloroplasts and a peripheral 1 - 3 layered chlorenchyma with elaborate cytoplasmic compartmentation. Each cell of the latter contains a peripheral cytoplasmic layer with scattered chloroplasts and a large globular cytoplasmic body located in the central vacuole which is densely packed with starch producing chloroplasts, and joined by the nucleus. Comparative studies on different leaves of the same individuals of ordinary C₄ species demonstrated a remarkable range of variation, from 1.0 ‰ in Salsola tragus up to 2.5 ‰ in Petrosimonia sibirica. This corresponds well with the variance in data from different individuals and populations. For explaining the larger interspecific variation from about 23 other species of Salsoloideae, the hypothesis is developed that in species or groups of taxa specific leakages of their C₄ systems do occur. By three characters unique in tribe Suaedeae and detected during the study, Bienertia is confirmed as well separated from Suaeda and as a monotypic genus.

Key words

Bienertia - Borszczowia - Chenopodiaceae - C₄ plants - facultative C₄/C₃ plants - C₃/C₄ intermediates - Kranz anatomy - carbon isotope ratio - δ¹³C values

1 The paper is dedicated to the memory of Vladimir Pyankov, the eminent student of C₄ Plant Biology, who tragically died on February 1st of 2002 during a research stay in Munich.
The paper is an extended version of a lecture presented at the 15th International Symposium “Biodiversity and Evolutionary Biology” in Bochum, Germany, 27. 9. 2001.

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1 The paper is dedicated to the memory of Vladimir Pyankov, the eminent student of C₄ Plant Biology, who tragically died on February 1st of 2002 during a research stay in Munich.
The paper is an extended version of a lecture presented at the 15th International Symposium “Biodiversity and Evolutionary Biology” in Bochum, Germany, 27. 9. 2001.

2 For discussion of a most questionable second species see p. 129.

3 For linguistic reasons, here we use the term borszczowioid instead of the earlier borszczovoid.

4 The terms austrobassioid and Kranz-suaedoid applied by Carolin et al. (1973) to leaves of Suaeda species are used here with hesitation. A revised terminology that covers more appropriately the diversity of leaves in Suaedeae is under study.

Section Editor: C. B. Osmond