RESUMO
The Portulacaceae enable the study of the evolutionary relationship between C4 and crassulacean acid metabolism (CAM) photosynthesis. Shoots of well-watered plants of the C3-C4 intermediate species Portulaca cryptopetala Speg. exhibit net uptake of CO2 solely during the light. CO2 fixation is primarily via the C3 pathway as indicated by a strong stimulation of CO2 uptake when shoots were provided with air containing 2% O2. When plants were subjected to water stress, daytime CO2 uptake was reduced and CAM-type net CO2 uptake in the dark occurred. This was accompanied by nocturnal accumulation of acid in both leaves and stems, also a defining characteristic of CAM. Following rewatering, net CO2 uptake in the dark ceased in shoots, as did nocturnal acidification of the leaves and stems. With this unequivocal demonstration of stress-related reversible, i.e. facultative, induction of CAM, P. cryptopetala becomes the first C3-C4 intermediate species reported to exhibit CAM. Portulaca molokiniensis Hobdy, a C4 species, also exhibited CAM only when subjected to water stress. Facultative CAM has now been demonstrated in all investigated species of Portulaca, which are well sampled from across the phylogeny. This strongly suggests that in Portulaca, a lineage in which species engage predominately in C4 photosynthesis, facultative CAM is ancestral to C4. In a broader context, it has now been demonstrated that CAM can co-exist in leaves that exhibit any of the other types of photosynthesis known in terrestrial plants: C3, C4 and C3-C4 intermediate.
Assuntos
Carbono/metabolismo , Ácidos Carboxílicos/metabolismo , Dióxido de Carbono/metabolismo , Luz , Filogenia , Folhas de Planta/metabolismo , Folhas de Planta/efeitos da radiação , Brotos de Planta/metabolismo , Brotos de Planta/efeitos da radiaçãoRESUMO
This study investigated whether Euphorbia subgenus Chamaesyce subsection Acutae contains C(3)-C(4) intermediate species utilizing C(2) photosynthesis, the process where photorespired CO(2) is concentrated into bundle sheath cells. Euphorbia species in subgenus Chamaesyce are generally C(4), but three species in subsection Acutae (E. acuta, E. angusta, and E. johnstonii) have C(3) isotopic ratios. Phylogenetically, subsection Acutae branches between basal C(3) clades within Euphorbia and the C(4) clade in subgenus Chamaesyce. Euphorbia angusta is C(3), as indicated by a photosynthetic CO(2) compensation point (Ð) of 69 µmol mol(-1) at 30 °C, a lack of Kranz anatomy, and the occurrence of glycine decarboxylase in mesophyll tissues. Euphorbia acuta utilizes C(2) photosynthesis, as indicated by a Ð of 33 µmol mol(-1) at 30 °C, Kranz-like anatomy with mitochondria restricted to the centripetal (inner) wall of the bundle sheath cells, and localization of glycine decarboxlyase to bundle sheath mitochondria. Low activities of PEP carboxylase, NADP malic enzyme, and NAD malic enzyme demonstrated no C(4) cycle activity occurs in E. acuta thereby classifying it as a Type I C(3)-C(4) intermediate. Kranz-like anatomy in E. johnstonii indicates it also utilizes C(2) photosynthesis. Given the phylogenetically intermediate position of E. acuta and E. johnstonii, these results support the hypothesis that C(2) photosynthesis is an evolutionary intermediate condition between C(3) and C(4) photosynthesis.
Assuntos
Euphorbia/fisiologia , Fotossíntese/fisiologia , Evolução Biológica , Dióxido de Carbono/análise , Dióxido de Carbono/metabolismo , Isótopos de Carbono/análise , Região do Caribe , Respiração Celular/fisiologia , Cloroplastos/ultraestrutura , Euphorbia/enzimologia , Euphorbia/ultraestrutura , Malato Desidrogenase/metabolismo , México , Mitocôndrias/ultraestrutura , Fosfoenolpiruvato Carboxilase/metabolismo , Filogenia , Folhas de Planta/enzimologia , Folhas de Planta/fisiologia , Folhas de Planta/ultraestrutura , Transpiração Vegetal/fisiologia , Ribulose-Bifosfato Carboxilase/metabolismo , Temperatura , TexasRESUMO
Using isotopic screens, phylogenetic assessments, and 45 years of physiological data, it is now possible to identify most of the evolutionary lineages expressing the C(4) photosynthetic pathway. Here, 62 recognizable lineages of C(4) photosynthesis are listed. Thirty-six lineages (60%) occur in the eudicots. Monocots account for 26 lineages, with a minimum of 18 lineages being present in the grass family and six in the sedge family. Species exhibiting the C(3)-C(4) intermediate type of photosynthesis correspond to 21 lineages. Of these, 9 are not immediately associated with any C(4) lineage, indicating that they did not share common C(3)-C(4) ancestors with C(4) species and are instead an independent line. The geographic centre of origin for 47 of the lineages could be estimated. These centres tend to cluster in areas corresponding to what are now arid to semi-arid regions of southwestern North America, south-central South America, central Asia, northeastern and southern Africa, and inland Australia. With 62 independent lineages, C(4) photosynthesis has to be considered one of the most convergent of the complex evolutionary phenomena on planet Earth, and is thus an outstanding system to study the mechanisms of evolutionary adaptation.