Origin and evolution of C4 photosynthesis in the tribe Salsoleae (Chenopodiaceae) based on anatomical and biochemical types in leaves and cotyledons

 The Chenopodiaceae genus Salsola contains a large number of species with C₄ photosynthesis. Along with derivative genera they have a prominent position among the desert vegetation of Asia and Africa. About 130 species from Asia and Africa were investigated to determine the occurrence of C₃ versus C₄ syndrome in leaves and cotyledons, and to study specific anatomical and biochemical features of photosynthesis in both photosynthetic organs. The species studied belong to all six previously identified sections of the tribe Salsoleae based on morphological characters. Types of photosynthesis were identified using carbon ¹³C/¹²C isotope fractionation. The representatives of all systematic groups were investigated for mesophyll anatomy and biochemical subtypes by determination of enzyme activity (RUBPC, PEPC, NAD- and NADP-ME and AAT) and primary photosynthetic products. Two photosynthetic types (C₃ and C₄) and two biochemical subtypes (NAD- and NADP-ME) were identified in both leaves and cotyledons. Both Kranz and non-Kranz type anatomy were found in leaves and cotyledons, but cotyledons had more diversity in anatomical structure. Strong relationships between anatomical types and biochemical subtypes in leaves and cotyledons were shown. We found convincing evidence for a similar pattern of structural and biochemical features of photosynthesis in leaves and cotyledons within systematic groups, and evaluated their relevance at the evolutionary level. We identified six groups in tribe Salsoleae with respect to photosynthetic types and mesophyll structure in leaves and cotyledons. Two separate lineages of biochemical and anatomical evolution within Salsoleae were demonstrated based on studies of leaves and cotyledons. The sections Caroxylon, Malpighipila, Cardiandra and Belanthera have no C₃ species and only the NAD-ME C₄ subtype has been found in leaves. We suggest the C₄ species in the NADP-ME lineage evolved in Coccosalsola and Salsola sections, and originated in the subsection Arbuscula. Coccosalsola contains many species with C₃ and/or C₃-C₄ intermediate photosynthesis. Within these main evolutionary lineages, species of different taxonomic groups (sections and subsections) had differences in anatomical or/and biochemical features in leaves and cotyledons. We conclude that structural and biochemical changes in the photosynthetic apparatus in species of the tribe Salsoleae were a key factor in their evolution and broad distribution in extreme desert environments. Received January 25, 2001 Accepted July 17, 2001     

A Contribution to the Life History of the Rare Indonesian Holoparasite Rafflesia patma (Rafflesiaceae)1

Fates and growth of Rafflesia patma buds and insect visitation to the flowers were monitored in an evergreen tropical rain forest in the Pangandaran Nature Reserve, Java, Indonesia, Of 59 buds marked in December 1985, 44 percent died before flowering, 7 percent flowered, and 49 percent were still in bud in May 1986. Mammals and birds caused the deaths of buds before they reached maturity. Only four buds flowered between December 1985 and May 1986; all were male. Rate of increase in diameter (cm/d) was much faster in large than in small buds. Once opening began, the flower opened fully in 24 to 48 hours; the flower remained open for three to five days before rotting. Only two genera of insects visited a male flower of R. patma: Lucilia and Sarcophaga (Diptera). Insect visitation was highest in the afternoon (1300–1700 h) and on the second or third day after flower opening. The increased number of flies visiting the flower coincided with the increasing odor emitted from it. Results of the present study were combined with information from the literature to construct a life cycle diagram of R. patma.