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     

Leaf and root growth dynamics in Eucalyptus globulus seedlings grown in drying soil

Summary Seedlings of Eucalyptus globulus growing in soil columns were subjected to a 24 day soil drying treatment. Water and solute potentials of both young expanding and fully expanded leaves declined under reduced soil water availability, while slightly higher turgor was sustained by the fully expanded leaves. Although leaf area of unwatered seedlings was smaller, the corresponding leaf dry weight was quite similar to that of well-watered seedlings. Soon after rewatering, leaf area of plants experiencing water shortage was comparable to that of well-watered plants. It seems that a difference in wall properties between juvenile and mature leaves allows for an effective pattern of water use by eucalypt plants growing in drying soil. Some stomatal opening is sustained and therefore, presumably, some carbon may be fixed, keeping the carbon balance of the whole plant positive, and allowing a continuous cell division despite the limited water supply. The highest root density of both well-watered and unwatered plants was found in the upper soil layers. However, root growth of unwatered seedlings was gradually increased in the deeper soil layers, where thicker root apices and higher soil water depletion rates per unit root length were recorded. As a consequence, root absorbing surface area was as large in unwatered plants as in well-watered plants.     

Allometric neoteny and the evolution of succulence in cacti

With the objective of analysing the role of heterochrony in the evolution of succulence in the cactus family, a comparative study of xylem development in six species with contrasting morphologies was carried out. Two woody leaf-bearing cacti and four succulent cactus species belong to different subdivisions within the family were analysed. In each species and for different ages, vessel-element length was measured, vessel-element lateral wall-pitting described and the percentage of xylem and parenchyma in the stem quantified. In the succulent species it was found that vessel element length did not change between juvenile and adult wood, that wall-pitting in adult plants was similar to that of seedlings, and that the woody tissue in adult plants was organized in vascular bundles as in the primary tissue of seedlingS. Leaf-bearing cacti, in contrast, changed in both vessel element length and wall-pitting when secondary wood was produced, and the secondary woody tissue of adult plants was organized in a continuous cambial cylinder as in most dicotyledonS. An allometric analysis suggests that a retardation in the developmental rate of woody tissues (allometric neoteny) is the main mechanism in the development of succulence in cacti.     

THE MUSCULAR HYDROSTAT OF THE FLORIDA MANATEE (TRICHECHUS MANATUS LATIROSTRIS): A FUNCTIONAL MORPHOLOGICAL MODEL OF PERIORAL BRISTLE USE

Facial musculature was examined in the Florida manatee, Tricbecbus manatus latirostris , in order to develop a functional model of perioral bristle use. Muscles identified include the M. levator nasolabialis, M. buccinatorius, M. maxillonasolabialis, M. centralis nasi, M. lateralis nasi, M. spbincter colli profundus pars oris, M. orbicularis oris, M. mandibularis, and M. mentalis. A new muscle, M. centralis nasi , has been named and is an integral part of perioral bristle movement. The snout of the Florida manatee is capable of performing complex movements. The prehensile ability of Florida manatees can be explained in the context of a muscular hydrostat as defined by Kier and Smith (1985). Eversion of certain bristles in the upper lip occurs by shortening longitudinal, transverse, and semicircular muscles in combination with volume displacement due to compensatory changes in the shape of the snout. Midline sweeping of these bristles is accomplished by the contraction of M. centralis nasi. Eversion of bristles on the lower jaw is a result of shortening of M. mentalis. Contraction of M. orbicularis oris pushes vegetation into the oral cavity. All observed movement patterns and uses of perioral bristles can be explained by variation of these sequences within the context of muscular hydrostat function.     

Vacchi's palatal organ: a widespread trait in Holocephali

A palatal organ, possibly used for food sorting and processing, has previously been identified among the vomerine toothplates of the chimaeroid Chimaera monstrosa. In this study, the palatal organ was described in six additional species, confirming it is a widespread trait among holocephalans. It is proposed that this palatal structure, which appears to differ in shape according to each chimaeroid's degree of durophagy and is not homologous to the palatal structure described in teleosts, be hereby referred to as Vacchi's organ.     

On the anatomy and taxonomy of Cerebratulus hepaticus Hubrecht, 1879 (Nemertini) from the Mediterranean (Banyuls-sur-Mer)

The anatomy of an insufficiently described species of heteronemertine, Cerebratulus hepaticus, is redescribed. The species is characterized by, inter alia, a cephalic lacuna with strands of longitudinal muscle fibres, a proboscis with three muscle layers, neurochord cells in the brain, a dermis in the foregut region with no or few longitudinal muscle fibres which is separated from the outer longitudinal muscle layer by connective tissue. The taxon is compared with two other Cerebratulus species, C. fuscus and C. marginatus. The material was collected at Banyuls-sur-Mer (France) in the Mediterranean.     

Leaf anatomy of Bruniaceae: ecological, systematic and phylogenetic aspects

CARLQUIST, S., 1991. Leaf anatomy of Bruniaceae: ecological, systematic and phylogenetic aspects. Quantitative and qualitative data are given for 60 species of the 12 genera of Bruniaceae; most data are based on liquid-preserved material. Leaves of Bruniaceae are basically linear (broader forms are probably derived) with an apicula that contains phellogen activity. Most bruniaceous leaves have some degree of isolateral construction, with transition to normal bifacial construction in a few species, but more commonly transition to 'inverse' bifacial structure (stomata on adaxial face, palisade on abaxial face). The latter type is correlated with the tendency for leaves to be appressed to stems. Tannins and very likely other dark-staining materials are very characteristic of mesophyll cells. Six genera have a large strand of fibres on the midvein and rhomboidal crystals in bundle sheath cells. The other six genera have few or no fibres on veins and have druses in mesophyll cells (but not in bundle sheath cells). These distinctions may relate to intrafamilial taxonomy, but they also support the primitive position usually accorded to Audouinia, Thamnea and Tittmannia. A key to genera based on leaf antomy is offered. Details of epidermal cell shape, cuticular relief and trichome form and structure based on scanning electron microscopy are given. Leaf anatomy, combined with other features, favours a relationship between Bruniaceae and Grubbiaceae in particular and in broader contexts allies Bruniaceae to rosalean and possibly hamamelidalean families.     

Evolution of a colour pattern: history,development, and selection

Patterns of melanin pigmentation in birds are extremely varied. Nevertheless it is easy to think of many patterns that are never observed, and others that frequently recur in diverse and distantly related species. Using as our model the avian genus Phylloscopus we ask how the restricted range of observed patterns might be attributable to a restricted range of variants produced by developmental perturbations. The patterns we consider consist of unmelanized patches on the wings, crown and rump on otherwise pigmented upperparts. We use reaction-diffusion models to show that gross features of the pattern can be simply predicted from considerations of embryo shape. We suggest that birds are expected to have more patterned heads, because the head region is relatively larger than other regions in the developing embryo. A comparative analysis across many species of birds and a phylogenetic analysis within the genus Phylloscopus show that the component elements of the pattern have repeatedly been lost and gained during evolution. A shift in a threshold reading could explain the appearance and disappearance of the unmelanized patches, perhaps through changes in the sensitivity of melanocytes to epidermal signals. Such threshold shifts would make the transition between patterned and unpatterned forms particularly easy once the patterns have been exposed to selection in some distant ancestor. This partitioning of the roles of selection and development implies that many features of the patterns reflect developmental mechanisms in both immediate and more distant ancestors.