scholarly journals The C4-pathway of photosynthesis. Evidence for an intermediate pool of carbon dioxide and the identity of the donor C4-dicarboxylic acid

1971 ◽  
Vol 125 (2) ◽  
pp. 425-432 ◽  
Author(s):  
M. D. Hatch

1. Leaves were exposed to 14CO2 under steady-state conditions for photosynthesis. The kinetics of entry or loss of label in pools of CO2 and other compounds was examined during the period of the pulse and a ‘chase’ with 12CO2. 2. With maize the kinetics of labelling of the major CO2 pool and of depletion of label during a ‘chase’ was consistent with this pool being derived from the C-4 of malate and being the precursor of the C-1 of 3-phosphoglycerate. 3. Similar results were obtained for Amaranthus leaves except that the C-4 of aspartate rather than malate was apparently the primary source of CO2. 4. The size and turnover time of the CO2 and C4 acid pools was calculated. These results provided the basis for estimating the concentration of CO2 in the bundle-sheath cells or chloroplasts assuming the pool was largely restricted to one or other of these compartments. 5. These findings are considered in relation to current schemes for the C4-pathway and the operation of a CO2 concentrating mechanism to serve ribulose diphosphate carboxylase.

1969 ◽  
Vol 47 (1) ◽  
pp. 15-21 ◽  
Author(s):  
T. Bisalputra ◽  
W. J. S. Downton ◽  
E. B. Tregunna

The ultrastructure of the chlorenchymatous tissues around the vascular bundles of three different types of grass leaves is described. In the temperate grass leaf, as exemplified by wheat, the inner mestom sheath contains proplastids. Normal chloroplasts are found only within the mesophyll cells. Smaller chloroplasts occur in cells of the ill-defined parenchymatic bundle sheath. This type of leaf has the photosynthetic pathway described by Calvin and a high carbon dioxide compensation value. In the tropical grasses, Sorghum and Aristida, the new photosynthetic pathway proposed by Hatch et al. and low carbon dioxide compensation are correlated with development of the parenchymatic bundle sheath. Cytological evidence indicates that cells of the bundle sheath are much more active than the surrounding mesophyll tissue. The specialized chloroplasts of the bundle sheath cells may be responsible for the physiological and biochemical differences between leaves of tropical and temperate grasses.


1969 ◽  
Vol 114 (1) ◽  
pp. 127-134 ◽  
Author(s):  
Hilary S. Johnson ◽  
M. D. Hatch

1. When leaves with the C4-dicarboxylic acid pathway of photosynthesis are exposed to 14CO2 the major labelled compounds formed, in order of labelling, are dicarboxylic acids, 3-phosphoglycerate, bexose phosphates and sucrose. During the present studies several quantitatively minor intermediates were identified and their labelling behaviour is described. 2. The pattern of labelling of dihydroxyacetone phosphate, fructose 1,6-diphosphate and ribulose di- and mono-phosphates during radiotracer pulse–chase experiments was consistent with their operation as intermediates in the pathway of carbon dioxide fixation. 3. Serine, glycine, alanine and glutamate had labelling patterns typical of products secondary to the main flow of carbon. 4. The mechanism of the transfer of label from C-4 of dicarboxylic acids to C-1 of 3-phosphoglycerate was also examined. Evidence consistent with pyruvate being derived from C-1, C-2 and C-3 of oxaloacetate, and for a relationship between ribulose 1,5-diphosphate and the acceptor for the C-4 carboxyl group, was obtained. 5. Evidence is provided that, under steady-state conditions, essentially all the label incorporated from 14CO2 into C-1 of 3 phosphoglycerate enters via C-4 of the dicarboxylic acids. These and other studies indicated that the route via dicarboxylic acids is essentially the sole route for entry of carbon into 3-phosphoglycerate.


1975 ◽  
Vol 2 (2) ◽  
pp. 111 ◽  
Author(s):  
MD Hatch ◽  
T Kagawa ◽  
S Craig

A selection of C4 species was surveyed to determine the relationship between their content of C4 acid decarboxylating enzymes, the activities of several other enzymes implicated in the C4 pathway, and their anatomical and ultrastructural features. The species examined clearly fell into three groups according to whether they contained high levels of either NADP malic enzyme (EC 1.1.1.40), phosphoenolpyruvate carboxykinase (EC 4.1.1.49) or NAD malic enzyme (EC 1.1 .1.39). The occurrence of high NADP malic enzyme activity was always associated with higher NADP malate dehydrogenase activity, while those species distinguished by high activities of either of the other two decarboxylases invariably contained high aspartate aminotransferase and alanine amino- transferase activities. Each of these decarboxylating enzymes was located in bundle sheath cells. NAD malic enzyme, but not phosphoenolpyruvate carboxykinase, was associated with mitochondria. Light and electron micrographs revealed differences between these groups with respect to the intracellular location of chloroplasts and mitochondria in bundle sheath cells, and the content and ultrastructure of mitochondria. The trend was for species with high NAD malic enzyme to contain the most mitochondria in the bundle sheath cells with apparently the most extensively developed cristae membrane systems. However, mitochondrial respiratory enzyme activities were similar for the three groups of species. The basic similarities and differences between the three groups of C4 plants distinguished by their differing C4 acid decarboxylating systems are discussed, and schemes for the probable photosynthetic reactions in bundle sheath cells are presented. A nomenclature to distinguish between these groups is proposed.


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