SummaryThe C4 photosynthetic pathway accounts for ∼25% of primary productivity on the planet despite being used by only 3% of species. Because C4 plants are higher yielding than C3 plants, efforts are underway to introduce the C4 pathway into the C3 crop rice. This is an ambitious endeavor; however, the C4 pathway evolved from C3 on multiple independent occasions over the last 30 million years, and steps along the trajectory are evident in extant species. One approach toward engineering C4 rice is to recapitulate this trajectory, one of the first steps of which was a change in leaf anatomy. The transition from C3 to so-called “proto-Kranz” anatomy requires an increase in organelle volume in sheath cells surrounding leaf veins. Here we induced chloroplast and mitochondrial development in rice vascular sheath cells through constitutive expression of maize GOLDEN2-LIKE genes. Increased organelle volume was accompanied by the accumulation of photosynthetic enzymes and by increased intercellular connections. This suite of traits reflects that seen in “proto-Kranz” species, and, as such, a key step toward engineering C4 rice has been achieved.

In this review, we examine how the specialized "Kranz" anatomy of C4 photosynthesis evolved from C3 ancestors. Kranz anatomy refers to the wreath-like structural traits that compartmentalize the biochemistry of C4 photosynthesis and enables the concentration of CO2 around Rubisco. A simplified version of Kranz anatomy is also present in the species that utilize C2 photosynthesis, where a photorespiratory glycine shuttle concentrates CO2 into an inner bundle-sheath-like compartment surrounding the vascular tissue. C2 Kranz is considered to be an intermediate stage in the evolutionary development of C4 Kranz, based on the intermediate branching position of C2 species in 14 evolutionary lineages of C4 photosynthesis. In the best-supported model of C4 evolution, Kranz anatomy in C2 species evolved from C3 ancestors with enlarged bundle sheath cells and high vein density. Four independent lineages have been identified where C3 sister species of C2 plants exhibit an increase in organelle numbers in the bundle sheath and enlarged bundle sheath cells. Notably, in all of these species, there is a pronounced shift of mitochondria to the inner bundle sheath wall, forming an incipient version of the C2 type of Kranz anatomy. This incipient version of C2 Kranz anatomy is termed proto-Kranz, and is proposed to scavenge photorespiratory CO2. By doing so, it may provide fitness benefits in hot environments, and thus represent a critical first stage of the evolution of both the C2 and C4 forms of Kranz anatomy.

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This study aims at proposing a new bioclimatic zonation for Iran based on the recently developed Global Bioclimatic Classification System (GBC) and tries to re-appraise this classification system by checking the degree of correspondence between its bioclimatic zones and the distribution of major phytogeographical regions, biomes, and a selection of plant and insect taxa of Iran. After application of the GBC to Iranian meteorological data, three macrobioclimates, ten bioclimates, and three bioclimatic variants were distinguished. The Mediterranean macrobioclimate is the dominant macrobioclimate and correlates with the Irano-Turanian biogeographical region, the Tropical macrobioclimate in southern Iran correlates with the Saharo-Sindian region and finally a small Temperate macrobioclimatic zone in northern Iran correlates with the Euro-Siberian region. Mediterranean macrobioclimate of Iran displays a wide range of bioclimates including Mediterranean pluviseasonal-oceanic, Mediterranean xeric-oceanic, Mediterranean pluviseasonal-continental, Mediterranean xeric-continental, Mediterranean desertic-oceanic, and Mediterranean desertic-continental. Tropical macrobioclimate also shows a range of bioclimates including Tropical xeric, Tropical desertic, and Tropical hyperdesertic. Finally, Temperate macrobioclimate has only one bioclimate i. e. Temperate oceanic. In conclusion, bioclimatic zones identified using the GBC system correlate well with the main phytogeographical regions of Iran at macrobioclimatic level and with major biomes at bioclimatic level. An advantage of the GBC over other bioclimatic classification systems is the inclusion of both seasonal and annual variations in those climatic parameters which are significant for the growth and development of plant populations and communities. GBC distinguishes several types of Mediterranean climate regimes for Iran and helps to differentiate between two different desert climate types i. e. Mediterranean and Tropical deserts in arid parts of the country. Because the climate in the Irano-Turanian region clearly differs from the Mediterranean region in its degree of continentality and seasonality, we propose the term ‘ xero-estival-continental’ or ‘Irano-Turanian’ instead of ‘Mediterranean’ bioclimate when describing the climate of continental Middle East and Central Asia.

HighlightC2 photosynthesis in grasses is facilitated by organelle enrichment in tandem with enhanced levels of GDC in the carbon-concentrating cells consistent with changes in expression of a single GLDP gene.