Characterizing the physics of plant root gravitropism: A systems modeling approach

Root gravitropism is divided into three mechanisms; the gravity sensor, transduction, and differential growth. The gravitropic response has been imitated with various mathematical constructs, but a coherent model based on systems engineering concepts does not exist. The goal of this research is to create models of the gravitropic sensor and differential growth response that are consistent with actual physical characteristics of these mechanisms.; The study initially establishes that the amyloplasts within the central columella cells of maize are feasible gravity sensors; statoliths. Video-microscopy studies of live root cap sections are used to quantify the dynamics of the statoliths. Extensive MATLAB analysis of amyloplast sedimentation indicates that an actin network interferes with the free sedimentation of the statoliths. This interference is most significant in the central region of the cell and less significant near the periphery. This obstruction of actin creates a channeling behavior in amyloplasts sedimenting through the cell's central region. The amyloplasts also appear to exhibit cross-correlated motions. Cytochalasin D mediates both the channeling and correlated behaviors, confirming that the obstructive influence is actin-based. The video analysis produced a refined value for maize cytoplasmic viscosity.; Efforts to model the differential growth mechanism examined historical growth data from numerous researchers. RELEL (relative elemental elongation) growth data applied to a model set analogous to bi-metallic bending is used. Testing and analysis of the model highlights an extremely high sensitivity of curvature to all RELEL parameters. This sensitivity appears to be the reason for the significant differences between gravitropic responses within like species.; Newly observed gravitropic responses, along with historical data, are used to explore the gravitropic time response specifications as opposed to averaging individual time-curvature data into single responses. This approach highlights the significant disadvantages of time-averaging, low sampling rates, and a lack of frequency components being incorporated into the response. A single feedback “black box” model is created so that, along with the sensor and differential growth models, some inferences could be made about the elusive transduction mechanism. Numerous pieces of circumstantial evidence are found that indicate that the gravitropic mechanism is not a single-pathway system.