Journal of Experimental Botany, Vol 50, 1779-1795, Copyright © 1999 by Oxford University Press
E Voznesenskaya, V Franceschi, V Pyankov and G Edwards
Certain members of the family Chenopodiaceae are the dominant species of
the deserts of Central Asia; many of them are succulent halophytes which
exhibit C4-type CO2 fixation of the NAD- or NADP-ME (malic enzyme)
subgroup. In four C4 species of the tribe Salsoleae, the Salsoloid-type
Kranz anatomy in leaves or stems was studied in relation to the diversity
in anatomy which was found in cotyledons. Halocharis
gossypina, has C4 NAD-ME Salsoloid-type photosynthesis in leaves
and C3 photosynthesis in dorsoventral non-Kranz cotyledons;
Salsola laricina has C4 NAD-ME Salsoloid-type leaves
and C4 NAD-ME Atriplicoid-type cotyledons; Haloxylon
persicum, has C4 NADP-ME Salsoloid-type green stems and C3
isopalisade non-Kranz cotyledons; and S. richteri has
C4 NADP-ME Salsoloid-type leaves and cotyledons. Immunolocalization studies
on Rubisco showed strong labelling in bundle sheath cells of leaves and
cotyledons of organs having Kranz anatomy. The C4 pathway enzyme
phosphoenolpyruvate carboxylase was localized in mesophyll cells, while the
malic enzymes were localized in bundle sheath cells of Kranz-type tissue.
Immunolocalization by electron microscopy showed NAD-ME is in mitochondria
while NADP-ME is in chloroplasts of bundle sheath cells in the respective
C4 types. In some C4 organs, it was apparent that subepidermal cells and
water storage cells also contain some chloroplasts which have Rubisco,
store starch, and thus perform C3 photosynthesis. In non-Kranz cotyledons
of Halocharis gossypina and Haloxylon
persicum, Rubisco was found in chloroplasts of both palisade and
spongy mesophyll cells with the heaviest labelling in the layers of
palisade cells, whereas C4 pathway proteins were low or undetectable. The
pattern of starch accumulation correlated with the localization of Rubisco,
being highest in the bundle sheath cells and lowest in the mesophyll cells
of organs having Kranz anatomy. In NAD-ME-type Kranz organs (leaves and
cotyledons of S. laricina and leaves of H.
gossypina the granal index (length of appressed membranes as a
percentage of total length of all membranes) of bundle sheath chloroplasts
is 1.5 to 2.5 times higher than that of mesophyll chloroplasts. In
contrast, in the NADP-ME-type Kranz organs (S.
richteri leaves and cotyledons and H.
persicum stems) the granal index of mesophyll chloroplasts is
1.5 to 2.2 times that of the bundle sheath chloroplasts. The mechanism of
photosynthesis in these species is discussed in relation to structural
differences.Keywords: Immunolocalization,
photosynthetic enzymes, C4 plants, anatomy, chloroplasts, ultrastructure,
Chenopodiaceae
ARTICLES
Anatomy, chloroplast structure and compartmentation of enzymes relative to photosynthetic mechanisms in leaves and cotyledons of species in the tribe Salsoleae (Chenopodiaceae)
Department of Anatomy and Morphology, VL Komarov Botanical Institute of Russian Academy of Sciences, Prof. Popof Street 2, 19736 St Petersburg, Russia; Botany Department, Washington State University, Pullman, WA 99164-4238, USA; Department of Plant Physiology, Ural State University, Ekaterinburg 620083, Russia; Corresponding author; Fax: +001 509 335 3517; E-mail: edwardsg@mail.wsu.edu
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