Ecophysiological Studies on the Vegetation of Madagascar: A δ13C and δD Survey for Incidence of Crassulacean Acid Metabolism (Cam) Among Orchids From Montane Forests and Succulents from the Xerophytic Thorn-Bush

Abstract

The incidence of Crassulacean acid metabolism (CAM) in plants collected at various habitats in Madagascar was investigated by survey of carbon and hydrogen isotope ratios ((δ¹³C and δD values). In about 50% of the epiphytic orchids from evergreen higher and lower montane forests the δ¹³C values were indicative for CAM. The remainder of the species are presumably C₃ plants. In all samples of malagasy epiphytic leafless orchids comprising 9 species, the δ¹³C values suggest extreme CAM with CO₂ uptake proceeding entirely during the night. All terrestrial orchids collected in the lower montane forests obviously acquire external carbon by C₃-photosynthesis, whereas Lissochilus decaryi, a terrestrial orchid from the semi-arid south of Madagascar and various other species of this genus are CAM plants. This is the first report of CAM occurrence in sympodial terrestrial orchids.

Judged by the δ¹³C values, all succulents (mainly Didiereaceae, Euphorbiaceae, Crassulaceae and Asclepiadaceae) collected at the xerophytic thorn-bush of the semi-arid south perform pronounced CAM. Where it applies, our δ¹³C measurements in the thorn-bush succulents revealed values being practically identical with those found by K. Winter in samples of the same species collected at the same site nearly 10 years earlier. This shows extreme constancy over long duration of time in the mode of CAM performed by the succulents of the malagasy thorn-bush vegetation. Since the δ¹³C survey now comprises all 11 known species of the Didiereaceae, it is unequivocally clear that all members of this family are CAM plants. Most of the individuals of the species of the Didiereaceae grown in a glass-house had slightly more negative δ¹³C values compared with those grown at the natural stands suggesting some contribution of C₃-photosynthesis to carbon acquisition under the evaporatively less demanding glass-house conditions (and perhaps higher CO₂ concentrations in the gas phase).

Despite of the fact that the hydrogen isotope composition of meteoric waters depends to a large extent on the altitude and temperature-climate of the site where the concerned plants grow, it was found that in samples obtained in the cooler higher evergreen montane forest as well as in the warmer lower evergreen montane forest and the lowland thorn-bush of the hot, semiarid south of Madagascar the δ values found in the organic matter (δD^org) were in the same range (between about - 10‰ to about - 90‰). This suggests that in our case the hydrogen isotope compositions of the meteoric waters were of minor importance in bringing about the δD^org values found in the plants.     

Minimum sum edge colorings of multicycles

In the minimum sum edge coloring problem, we aim to assign natural numbers to edges of a graph, so that adjacent edges receive different numbers, and the sum of the numbers assigned to the edges is minimum. The chromatic edge strength of a graph is the minimum number of colors required in a minimum sum edge coloring of this graph. We study the case of multicycles, defined as cycles with parallel edges, and give a closed-form expression for the chromatic edge strength of a multicycle, thereby extending a theorem due to Berge. It is shown that the minimum sum can be achieved with a number of colors equal to the chromatic index. We also propose simple algorithms for finding a minimum sum edge coloring of a multicycle. Finally, these results are generalized to a large family of minimum cost coloring problems.     

On the growth of components with non-fixed excesses

Denote by an l-component a connected graph with l edges more than vertices. We prove that the expected number of creations of (l+1)-component, by means of adding a new edge to an l-component in a randomly growing graph with n vertices, tends to 1 as l,n tends to ∞ but with l=o(n^1/4). We also show, under the same conditions on l and n, that the expected number of vertices that ever belong to an l-component is (12l)^1/3n^2/3.