Date of Award

January 2016

Degree Type


Degree Name

Doctor of Philosophy (PhD)


Agricultural and Biological Engineering

First Advisor

David M Umulis

Committee Member 1

Doraiswami Ramkrishna

Committee Member 2

Gregory T Buzzard

Committee Member 3

Kevin V Solomon


Organs and tissue development often experience perturbations, but developmental processes seem to replicate a common body template to maintain appropriate proportions and positions. The key signaling factors that guide a number of those processes are known as morphogens. Developing cells sense their respective positional information from a graded morphogen profile, and differentiate into patterns. Remarkably, patterns are highly robust and reproducible among species, and the underlying mechanisms associated with such high degrees of precision are still enigmatic. In addition, details of the signal, such as the Bone Morphogenetic Protein (BMP) signal, that transmit patterning information to a group of homogenous cells to differentiate is not well understood. Determining how developmental processes ensure robust patterning in the presence of perturbations maintain structural precision by scaling, and what regulatory mechanisms act to ensure robust and reproducible patterning are two longstanding questions that need unraveling. Moreover, determining the mechanisms by which BMP homodimers dominate signaling in developing zebrafish embryos and other contexts is a key factor in understanding developmental regulation for a classic morphogen patterning.