Abstract
Experimental studies of the flowering of Arabidopsis thaliana have shown that a large complex gene regulatory network (GRN) is responsible for its regulation. This process has been mathematically modelled with deterministic differential equations by considering the interactions between gene activators and inhibitors (Valentim et al. in PLoS ONE 10(2):e0116973, 2015; van Mourik et al. in BMC Syst Biol 4(1):1, 2010). However, due to complexity of the model, the properties of the network and the roles of the individual genes cannot be deducted from the numerical solution the published work offers. Here, we propose simplifications of the model, based on decoupling of the original GRN to motifs, described with three and two differential equations. A stable solution of the original model is sought by linearisation of the original model which contributes to further investigation of the role of the individual genes to the flowering. Furthermore, we study the role of noise by introducing and investigating two types of stochastic elements into the model. The deterministic and stochastic nonlinear dynamic models of Arabidopsis flowering time are considered by following the deterministic delayed model introduced in Valentim et al. (2015). Steady-state regimes and stability of the deterministic original model are investigated analytically and numerically. By decoupling some concentrations, the system was reduced to emphasise the role played by the transcription factor Suppressor of Overexpression of Constants1 (textit{SOC}1) and the important floral meristem identity genes, Leafy (textit{LFY}) and Apetala1 (textit{AP}1). Two-dimensional motifs, based on the dynamics of textit{LFY} and textit{AP}1, are obtained from the reduced network and parameter ranges ensuring flowering are determined. Their stability analysis shows that textit{LFY} and textit{AP}1 are regulating each other for flowering, matching experimental findings. New sufficient conditions of mean square stability in the stochastic model are obtained using a stochastic Lyapunov approach. Our numerical simulations demonstrate that the reduced models of Arabidopsis flowering time, describing specific motifs of the GRN, can capture the essential behaviour of the full system and also introduce the conditions of flowering initiation. Additionally, they show that stochastic effects can change the behaviour of the stability region through a stability switch. This study thus contributes to a better understanding of the role of textit{LFY} and textit{AP}1 in Arabidopsis flowering.
Highlights
Arabidopsis thaliana is a small, annual flowering plant in the Brassicaceae family which is a favourite model organism for plant biology research due mainly to its small size, simple genome and rapid life cycle
We consider the deterministic dynamic model of delay differential equations (DDEs) describing the flowering of the Arabidopsis species proposed by Valentim et al (2015)
The model is based on a feedback loop, constructed with eight genes, where six of them are internal: Apetala1 (AP1), Leafy (LFY), Suppressor of Overexpression of Constants 1 (SOC1), Agamous-Like 24 (AGL24), Flowering Locus T (FT) and FD
Summary
Arabidopsis thaliana is a small, annual flowering plant in the Brassicaceae (mustard) family which is a favourite model organism for plant biology research due mainly to its small size, simple genome and rapid life cycle. Physiological and environmental conditions of the plant regulate the timing of transition for the optimal reproductive achievement, and their reactions are integrated into a complex GRN which monitors and regulates this transition (Kardailsky et al 1999; Levy and Dean 1998; Wellmer and Riechmann 2010) Genes and their regulatory interactions are significant factors in biological systems at the molecular level since the understanding of their impact on each other’s regulation is crucial to comprehend the response of gene disturbances on flowering time (Valentim et al 2015). Numerous genes appear to be acting as flowering time regulators of Arabidopsis thaliana (Ryan et al 2015), and different pathways have been constructed to reveal the flowering of this plant (Amasino 2010; Greenup et al 2009; Kardailsky et al 1999; Yant et al 2009) This complex network of many interacting genes can be dynamically modelled using systems with many equations (Jaeger et al 2013; Valentim et al 2015; van Mourik et al 2010; Wang et al 2014). The other two genes are considered as external inputs: Short Vegetative Phase (SVP) and Flowering Locus C (FLC)
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