Timing Is Everything： Highly Specific and Transient Expression of a MAP Kinase Determines Auxin-Induced Leaf Venation Patterns in Arabidopsis

Mitogen-activated protein kinase （MAPK） cascades are universal signal transduction modules present in all eukaryotes. In plants, MAPK cascades were shown to regulate cell division, developmental processes, stress responses, and hormone pathways. The subgroup A of Arabidopsis MAPKs consists of AtMPK3, AtMPK6, and AtMPK10. AtMPK3 and AtMPK6 are activated by their upstream MAP kinase kinases （MKKs） AtMKK4 and AtMKK5 in response to biotic and abiotic stress. In addition, they were identified as key regulators of stomatal development and patterning. AtMPKIO has long been considered as a pseudo-gene, derived from a gene duplication of AtMPK6. Here we show that AtMPKIO is expressed highly but very transiently in seedlings and at sites of local auxin maxima leaves. MPK10 encodes a functional kinase and interacts with the upstream MAP kinase kinase （MAPKK） AtMKK2. mpklO mutants are delayed in flowering in long-day conditions and in continuous light. Moreover, cotyledons of mpk10 and mkk2 mutants have reduced vein complexity, which can be reversed by inhibiting polar auxin transport （PAT）. Auxin does not affect AtMPKIO expression while treatment with the PAT inhibitor HFCA extends the expression in leaves and reverses the mpklO mutant phenotype. These results suggest that the AtMKK2-AtMPK10 MAPK module regulates venation complexity by altering PAT efficiency.

Plant ontogenetic changes in vein and stomatal traits and their relationship with economic traits in leaves of three Mediterranean oaks

Aims We compared vein and stomatal traits of seedlings and adults of three Mediterranean Quercus species.Previous work suggests that gas-exchange rates tend to be higher at the seedling stage than in adults.Our objective was to determine whether vein and stomatal traits vary throughout whole-plant ontogeny in parallel with the changes in gas-exchange rates.We addressed the following alternative hypotheses:hypothesis 1—seedlings show higher vein and stomatal densities than adults;and hypothesis 2—seedlings have lower investments in vascular tissues to reduce construction costs.Methods Ten specimens from each growth stage were randomly sampled for each species in a location in central-western Spain.We measured mean stomatal and vein traits(size and number of stomata per unit of leaf area,vein density,vein volume,vein to epidermis distance),leaf mass per area and lamina thickness.Important Findings Minor vein density and vein volume per area increased with tree age,which seems inconsistent with the ontogenetic trends in gas-exchange rates.This discrepancy is in support of our hypothesis 2,and it suggests that,at the seedling stage,reducing investments in vascular tissues in benefit of maximizing growth rates is a priority.Larger interveinal distances in seedlings were compensated by smaller vein to epidermis distances.The thin leaves of the seedlings may thus constitute as a necessary trait for achieving shorter path length distances for the transport of water to evaporation sites without the need for a strong investment in costly vascular tissues.