Mathematical modelling of photoperiodic external coincidence mechanisms in the model plant, Arabidopsis thaliana
Smith, Robert William
As plants are sessile organisms, processes controlling plant growth and development must react to fluctuations in the external environment to aid plant survival. However, as the climate of the Earth changes and becomes more extreme, plants become less able to develop to their optimal capacity and this can have an adverse effect on crop yield and biofuel feedstock production. Thus, it is becoming increasingly important to understand the molecular mechanisms used by plants to respond to external stimuli. One important system that plants utilise in their response to environmental fluctuations is the circadian clock. The circadian clock is a time-measuring device that buffers the timing of plant growth and development against fluctuations in the local environment, such as temperature, light quality and light intensity. Importantly, the circadian clock is also able to measure day-length (photoperiod). Thus, plant development and growth is co-ordinated with photoperiod that is closely linked to seasonal changes. A key example of this is the time taken for a plant to flower. Flowering of Arabidopsis thaliana occurs specifically in long-days (LDs) of spring/summer months. Thus, the circadian clock is a key regulator promoting flowering in LD conditions. In conjunction with experimental studies, mathematical modelling has proven to be a successful method of elucidating the mechanisms that underlie complex biological systems. One example of this 'systems biology' approach is in uncovering the components that make up the Arabidopsis circadian clock mechanism. Previous research in our group has also led to the development of a model describing photoperiodic flowering that is tentatively linked to the circadian clock mechanism. In this thesis I shall develop on these models to highlight five key results: 1. using rhythmic PHYTOCHROME INTERACTING FACTOR 4 (PIF4) and PIF5 mRNA as an example, I shall show that multiple circadian regulators are required to describe rhythmic transcription of target genes across multiple photoperiods; 2. the stabilisation of CONSTANS (CO) protein by the blue light-signalling component FLAVIN-BINDING, KELCH REPEAT, F-BOX 1 (FKF1) is required to for flowering in LDs and has a relatively larger impact on photoperiodic flowering than FKF1-dependent degradation of CYCLING DOF FACTOR 1 (CDF1), an inhibitor of flowering; 3. multiple components of the circadian clock play specific post-translational roles in photoperiodic flowering to promote the acceleration of flowering specifically in LDs; 4. temperature regulation of photoperiodic flowering can be explained through an interaction between CO and PIF proteins, limiting the effects of temperature to a specific time-window in a 24-hour day; 5. red light- and temperature-control of the circadian clock can be explained by altering the post-translational regulation of circadian clock components.