Plants are better engineers: the complexity of plant organ morphogenesis.

Plant are better engineers than us. They reproducibly create new organs while we are still far from being able to engineer plant morphogenesis. It is challenging to understand plant morphogenesis due to its complexity. Complex intersecting regulatory networks often mask general principles. Cells and molecular regulators typically behave variably yet the plant uses these inputs to achieve robust outcomes. Regulatory networks often act in the non-linear range near tipping points such that small stochastic variations are used to make important developmental decisions. With the recent employment of 4D growth tracking combined with quantitative analysis of regulatory networks and computational modeling, we now have better capacity to explore and embrace the complexity of plant organ morphogenesis.

Cytokinin and CLE signaling are highly intertwined developmental regulators across tissues and species.

The control of cell identity and differentiation is critical for proper development. In plants, cell identity is largely determined by a cell's spatial context, which is communicated in the form of varying abundances of hormones. Two classes of hormones, the classical phytohormone cytokinin and the small CLE peptide hormones, are potent regulators of cell division and cell differentiation. While a relationship between these two classes of hormones is well-established at developing shoot tips, recent evidence suggests that CLE and cytokinin signaling converge on the same developmental processes across many different contexts and in widely divergent species. Here, we review evidence predominately from Arabidopsis thaliana and the moss Physcomitrella patens that supports a general model where CLE and cytokinin signaling are highly intertwined developmental regulators with antagonistic functions in shoots and synergistic functions in roots.

Why plants make puzzle cells, and how their shape emerges.

The shape and function of plant cells are often highly interdependent. The puzzle-shaped cells that appear in the epidermis of many plants are a striking example of a complex cell shape, however their functional benefit has remained elusive. We propose that these intricate forms provide an effective strategy to reduce mechanical stress in the cell wall of the epidermis. When tissue-level growth is isotropic, we hypothesize that lobes emerge at the cellular level to prevent formation of large isodiametric cells that would bulge under the stress produced by turgor pressure. Data from various plant organs and species support the relationship between lobes and growth isotropy, which we test with mutants where growth direction is perturbed. Using simulation models we show that a mechanism actively regulating cellular stress plausibly reproduces the development of epidermal cell shape. Together, our results suggest that mechanical stress is a key driver of cell-shape morphogenesis.

Use it or average it: stochasticity in plant development.

A process that is stochastic has a probabilistic or randomly determined outcome. At the molecular level, all processes are stochastic; but development is highly reproducible, suggesting that plants and other multicellular organisms have evolved mechanisms to ensure robustness (achieving correct development despite stochastic and environmental perturbations). Mechanisms of robustness can be discovered through isolating mutants with increased variability in phenotype; such mutations do not necessarily change the average phenotype. Surprisingly, some developmental robustness mechanisms actually exploit stochasticity as a useful source of variation. For example, gene expression is stochastic and can be utilized to create subtle differences between identical cells that can initiate the patterning of specialized cell types. Stochasticity can also be used to promote robustness through spatiotemporal averaging-stochasticity can be averaged out across space and over time. Thus, organisms often harness stochasticity to ensure robust development.