Defining the mechanisms of specificity in the symbiosis signalling pathway of Medicago truncatula

Miller, Ben (2012) Defining the mechanisms of specificity in the symbiosis signalling pathway of Medicago truncatula. Doctoral thesis, University of East Anglia.

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    Legume plants are able to form symbiotic interactions with rhizobial bacteria and
    arbuscular mycorrhizal fungi. The establishment of these two symbioses depends
    upon signalling between the plant host and the microorganism, of which
    lipochitooligosaccharide (LCO) signals are essential. Perception of symbiotic LCOs
    induces a signalling pathway which is common to both mycorrhization and
    nodulation, and oscillations of calcium in the nucleus (so-called calcium spiking) are
    central to this common symbiosis signalling pathway. Detailed analysis of Medicago
    truncatula gene expression in response to rhizobial and mycorrhizal LCOs reveals
    relatively little overlap in gene induction. The nodulation-specific marker NIN was
    induced by both rhizobial and sulphated mycorrhizal LCOs. However, the
    mycorrhization-specific marker MSBP1 was only induced by non-sulphated
    mycorrhizal LCOs. Importantly, this differential induction of NIN and MSBP1 by
    LCOs was dependent on components of the common symbiosis signalling pathway.
    Immediately downstream of calcium spiking lies a calcium- and
    calcium/calmodulin-dependent protein kinase (CCaMK) which is believed to decode
    calcium spiking. It has been hypothesised that CCaMK activates differential
    signalling outputs in response to specific calcium signatures associated with each
    symbiosis. A model for the activation of CCaMK via autophosphorylation has
    previously been proposed, but is unable to explain symbiosis signalling specificity.
    A mutational approach was therefore undertaken to determine the importance of
    calcium- and calmodulin-binding during the activation of CCaMK. Interestingly,
    most mutations which blocked nodulation also blocked mycorrhization. However,
    the identification of a mutation which impairs mycorrhization but permits
    nodulation suggests that CCaMK may mediate specificity via this putative
    phosphorylation site.
    A multidisciplinary study based on this mutational analysis has proposed a new
    model for CCaMK activation which relies upon differentially regulated
    autophosphorylation of CCaMK in response to calcium and calmodulin. A full
    mechanism to explain how CCaMK determines specificity has yet to be elucidated.

    Item Type: Thesis (Doctoral)
    Faculty \ School: Faculty of Science > School of Biological Sciences
    Depositing User: Mia Reeves
    Date Deposited: 02 May 2013 14:19
    Last Modified: 02 May 2013 14:19

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