Cell polarity: Regulators and mechanisms in plants

Cell polarity plays an important role in a wide range of biological processes in plant growth and development.Cell polarity is manifested as the asymmetric distribution of molecules,for example,proteins and lipids,at the plasma membrane and inside of a cell.Here,we summarize a few polarized proteins that have been characterized in plants and we review recent advances towards understanding the molecular mechanism for them to polarize at the plasma membrane.Multiple mechanisms,including membrane trafficking,cytoskeletal activities,and protein phosphorylation,and so forth define the polarized plasma membrane domains.Recent discoveries suggest that the polar positioning of the proteo-lipid membrane domain may instruct the formation of polarity complexes in plants.In this review,we highlight the factors and regulators for their functions in establishing the membrane asymmetries in plant development.Furthermore,we discuss a few outstanding questions to be addressed to better understand the mechanisms by which cell polarity is regulated in plants.     

Cell polarity: fixing cell polarity with Pins

A protein complex is assembled in a step-wise manner at the apical pole of Drosophila neuroblasts. This complex organizes the apical-basal polarity of asymmetrically dividing neuroblasts, and may act via G-protein signalling.     

Cell polarity: formin on the move

Formins assemble actin filaments that are typically arranged in long bundles. A new study has discovered that a fission yeast polarity formin transiently assembles short actin filaments at the cell tip, and then releases from the cortex and rides into the cell interior on filaments within the bundle.     

Cell polarity: a tale of two Ts

The microtubule cytoskeleton plays an important role in cell polarity. Central to this process in fission yeast is tea1p, a marker of polarized cell growth that is delivered to the cell surface in a microtubule-dependent fashion. Recent studies suggest that the actin-binding protein bud6p may be a tea1p effector.     

Cell polarity

Despite extensive genetic analysis of the dynamic multi-phase process that transforms a small population of lateral plate mesoderm into the mature limb skeleton, the mechanisms by which signaling pathways regulate cellular behaviors to generate morphogenetic forces are not known. Recently, a series of papers have offered the intriguing possibility that regulated cell polarity fine-tunes the morphogenetic process via orienting cell axes, division planes and cell movements. Wnt5a-mediated non-canonical signaling, which may include planar cell polarity, has emerged as a common thread in the otherwise distinct signaling networks that regulate morphogenesis in each phase of limb development. These findings position the limb as a key model to elucidate how global tissue patterning pathways direct local differences in cell behavior that, in turn, generate growth and form.     

Rop GTPase: a master switch of cell polarity development in plants

Cell polarity is fundamentally important to plant growth and development, yet the mechanism governing its development is understood poorly. Several studies have revealed a role for Rop GTPases in pollen polar tip growth. Rop is also localized to the future site of root hair development and the tip of root hairs, and expression of constitutively active Rop mutants impacts on the morphogenesis of tip-growing root hairs as well as on non-tip-growing cells. These findings highlight the importance of Rop as a common switch in cell polarity control in plants.     

Cell polarity: squaring the circle

Recent studies have found that Drosophila gene products required for zonula adherens formation in the ectoderm are also involved in the asymmetric cell division of the neuroblast. The results illustrate the reiterated use of groups of proteins to dictate cell polarity in epithelial and non-epithelial cells.     

Cell polarity: the PARty expands

Recent work has revealed an evolutionarily conserved trio of proteins that regulate cell polarity in epithelial cells, embryonic blastomeres and neural precursors. This common cell-polarity mechanism is used in cell-specific ways, as highlighted by the recent finding that at least two different types of asymmetric division are observed in Drosophila neural precursors.     

Cell axiality and polarity in plants — adding pieces to the puzzle

Plant cell polarity is important for cellular function and multicellular development. Classical physiological and cell biological analyses identified cues that orient cell polarity and suggested molecules that translate a cue into intracellular asymmetry. A range of proteins that either mark or are involved in the establishment of a (polar) axis are now available, as are many relevant mutants. These tools are likely to facilitate a dissection of the molecular mechanisms behind cell and organ polarity in the near future.     

Cell polarity in plants: when two do the same, it is not the same...

In unicellular and multicellular organisms, cell polarity is essential for a wide range of biological processes. An important feature of cell polarity is the asymmetric distribution of proteins in or at the plasma membrane. In plants such polar localized proteins play various specific roles ranging from organizing cell morphogenesis, asymmetric cell division, pathogen defense, nutrient transport and establishment of hormone gradients for developmental patterning. Moreover, flexible respecification of cell polarities enables plants to adjust their physiology and development to environmental changes. Having evolved multicellularity independently and lacking major cell polarity mechanisms of animal cells, plants came up with alternative solutions to generate and respecify cell polarity as well as to regulate polar domains at the plasma membrane.