Abstract
Organ morphogenesis is the process of shape acquisition initiated with a small reservoir of undifferentiated cells. In plants, morphogenesis is a complex endeavor that comprises a large number of interacting elements, including mechanical stimuli, biochemical signaling, and genetic prerequisites. Because of the large body of data being produced by modern laboratories, solving this complexity requires the application of computational techniques and analyses. In the last two decades, computational models combined with wet-lab experiments have advanced our understanding of plant organ morphogenesis. Here, we provide a comprehensive review of the most important achievements in the field of computational plant morphodynamics. We present a brief history from the earliest attempts to describe plant forms using algorithmic pattern generation to the evolution of quantitative cell-based models fueled by increasing computational power. We then provide an overview of the most common types of “digital plant” paradigms, and demonstrate how models benefit from diverse techniques used to describe cell growth mechanics. Finally, we highlight the development of computational frameworks designed to resolve organ shape complexity through integration of mechanical, biochemical, and genetic cues into a quantitative standardized and user-friendly environment.
Highlights
Plant development has inspired the interest of scientific minds since antiquity
We provide an overview of the most common types of “digital plant” paradigms, and demonstrate how models benefit from diverse techniques used to describe cell growth mechanics
These formulas could not represent important mechanical aspects exhibited by living cells such as water transpiration, wall stress relaxation, pressure relaxation, and elastic deformations (Geitmann and Ortega, 2009), which were addressed by follow-up studies: Cosgrove (1986) extended Lockhart’s paradigm to multicellular organization; Silk and Wagner (1980) proposed a non-compartmented continuum model; Ortega (1985) augmented Lockhart’s equation with an elastic component, while Veytsman and Cosgrove (1998) rederived the formula in terms of thermodynamics of polymer networks
Summary
Plant development has inspired the interest of scientific minds since antiquity. The first attempt to formulate plant growth into a mathematically coherent framework was given by Thompson (1917) in his landmark book On Growth and Form. The relentless increase of computational power of modern machines allowed the definition of increasingly complex structures, such as tissues and entire organs (Glazier and Graner, 1993) These initial successes in modeling plant development motivated the incorporation of biomechanics properties to further increase the realism of plant form generation. Lockhart (1965) proposed a model to predict cell wall growth rate from internal turgor pressure by formulating a set of rules regulating the behavior of an idealized cell wall, which will later become the de facto standard for mechanical modeling of plant cell elongation These formulas could not represent important mechanical aspects exhibited by living cells such as water transpiration, wall stress relaxation, pressure relaxation, and elastic deformations (Geitmann and Ortega, 2009), which were addressed by follow-up studies: Cosgrove (1986) extended Lockhart’s paradigm (which only applied to single cells in isolation) to multicellular organization; Silk and Wagner (1980) proposed a non-compartmented continuum model; Ortega (1985) augmented Lockhart’s equation with an elastic component, while Veytsman and Cosgrove (1998) rederived the formula in terms of thermodynamics of polymer networks. It has been suggested that even isotropic growth could generate anisotropic patterns as a result
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