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75 Integrating Membrane Transport with Male Gametophyte ... - TAIR

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359 eQTL-mapping reveals AMP2A, a circadian regulated dioxygenase affecting rhythmic leaf<br />

movement<br />

Samuel Hazen 1 , Tatjana Singer 2 , Yiping Fan 3 , Steven Briggs 4 , Steve Kay 1<br />

1<br />

The Scripps Research Institute, 2 The Salk Institute, 3 St. Jude Children's Research Hospital, 4 University of<br />

California, San Diego<br />

Organisms have evolved to coordinate their activities <strong>with</strong> the day-night cycle caused by the Earth's rotation <strong>with</strong> an<br />

internal mechanism known as the circadian clock. Anticipating environmental changes rather than simply reacting to them<br />

increases photosynthesis, growth, and survival. Arabidopsis accessions often exhibit diverse circadian properties that are<br />

under genetic control. We combined high-density array genotyping and transcription profiling the Col/Ler recombinant<br />

inbred line population to explore this natural variation and identified an eQTL associated <strong>with</strong> leaf movement rhythms.<br />

The expression of a putative dioxygenase located in a previously mapped QTL was strongly correlated <strong>with</strong> the amplitude<br />

of leaf rhythms. We discovered that the Col allele of the putative causal gene is rhythmically expressed while the Ler<br />

allele, which has a positive effect on amplitude, is not expressed at all. The expression polymorphism occurs in at least<br />

one third of all accessions tested and is likely caused by a 23bp deletion in the upstream regulatory region. Moreover, a<br />

loss-of-function mutant in Col exhibited increased amplitude and thus confirmed the phenotype association. Flowering<br />

time and the expression of several core clock genes are not altered in the loss-of-function mutant; therefore, the putative<br />

dioxygenase appears to be part of circadian regulated output pathway.<br />

360 Arabidopsis/oomycete symbioses provide a model for phylogenomics and molecular<br />

epidemiology in plants<br />

Eric Holub<br />

University of Warwick<br />

Much progress has been made from molecular genetics of disease resistance, <strong>with</strong> Arabidopsis thaliana pathology at<br />

the forefront in revealing parallels <strong>with</strong> innate immunity of animals. Since coevolution emerged as a research topic more<br />

than 25 years ago, there has been general agreement that disease resistance evolves <strong>with</strong>in plant species by a treadmill of<br />

balancing selection involving parasite recognition proteins encoded by R-genes and matching “avirulence” effectors (AVR<br />

genes) in the parasite (1). Evidence for an adaptive “arms race” driving speciation of the host as well as the parasite has<br />

yet to be revealed. This may begin to change <strong>with</strong> phylogenetic analyses of host genes that govern basic compatibility in<br />

plant-parasite symbioses, or that delimit host specialization of parasite taxa. Ellingboe (2) defined basic compatibility as<br />

an evolved state that exists between taxa (species or subspecies) of a plant and a parasite that has adapted to survive and<br />

proliferate in that host. The adapted parasite must be equipped to evade or overcome constitutive barriers and inducible<br />

defenses of its compatible host, and possess the developmental and metabolic characteristics necessary to exploit its nutritious<br />

environment. He suggested that the evolution of R-AVR mediated resistance is superimposed onto basic compatibility.<br />

The parasite secretome of effector-like genes and any known corresponding host R-genes will provide a focus for future<br />

research. Genome-wide comparative biology among species of hosts and parasites would constitute phylogenomics of<br />

parasitic symbioses, whereas variability <strong>with</strong>in species would enable microevolutionary or molecular epidemiology. Research<br />

of Arabidopsis associations <strong>with</strong> its adapted oomycete parasites (Hyaloperonospora parasitica and Albugo candida) will<br />

provide pull-through knowledge into crops and other wild plant species (1). Exciting progress has recently been made in<br />

molecular characterisation of effector proteins from H. parasitica, based on recognition by corresponding Arabidopsis R-<br />

genes. A large number of effector-like proteins may be gleaned and classified by bioinformatics from a reference genome<br />

of H. parasitica (isolate HpEmoy2) and developed for different experimental uses in comparative biology. Some may prove<br />

useful for phylogenomics among species, and others may prove most useful for molecular epidemiology.<br />

1 Holub, EB. Curr. Opin. Plant Biol. 2006 in press.<br />

2 Ellingboe, AH. In Physiological Plant Pathology (Encycl. of Plant Physiol., Vol 4). Heitefuss R, Williams, PH (eds): Springer. 1976:761-778.

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