Bacterial cell polarity: a "swarmer-stalked" tale of actin

The actin cytoskeleton is important for cell polarity and morphogenesis in eukaryotic organisms. A recent article describes an unexpected requirement for the actin-like protein MreB in the polarization of the bacterium Caulobacter crescentus. More surprisingly, the formation of a filamentous MreB structure that traverses the length of the cell is sufficient for randomized polar localization of cell-fate proteins. In this article, we discuss the significance of these findings and the possible mechanisms by which an actin-like cytoskeleton could mediate cell polarity in bacteria.     

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.     

Recycling polarity

Three current papers in Cell and in this issue of Developmental Cell highlight the role of the exocyst in recycling of membrane proteins from endosomes to the plasma membrane in asymmetric cell division and polarized epithelial cells.     

Planar polarity

The clinical burden of both adult and neonatal lung disease worldwide is substantial; in the UK alone, respiratory disease kills one in four people. It is increasingly recognized that genes and pathways that regulate lung development, may be aberrantly activated in disease and/or reactivated as part of the lungs’ intrinsic repair mechanisms. Investigating the genes and signaling pathways that regulate lung growth has led to significant insights into the pathogenesis of congenital and adult lung disease. Recently, the planar cell polarity (PCP) pathway has been shown to be required for normal lung development, and data suggests that this signaling pathway is also involved in the pathogenesis of some lung diseases. In this review, we summarize current evidence indicating that the PCP pathway is required for both lung development and disease.     

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.     

Axonal polarity in Drosophila wings with mutant cuticular polarity patterns

In Drosophila melanogaster certain mutations alter the polarity of trichomes and bristles, cuticular structures secreted by the epithelial cells of the adult fly. Since sensory neurons arise from epithelial cell precursors, and sensory axons grow along the inner faces of epithelial cells, we have studied the developing wings of these mutants to see whether the change in epithelial cell polarity has an influence on the direction of axon outgrowth. The nerve patterns formed in the mutants prickled, inturned, and frizzled, however, were largely normal, indicating that in these cases the polarity of the cuticular structures produced by the epithelial cells is altered without any effect on the polarity of the associated axons.     

Mitotic internalization of planar cell polarity proteins preserves tissue polarity

Planar cell polarity (PCP) is the collective polarization of cells along the epithelial plane, a process best understood in the terminally differentiated Drosophila wing. Proliferative tissues such as mammalian skin also show PCP, but the mechanisms that preserve tissue polarity during proliferation are not understood. During mitosis, asymmetrically distributed PCP components risk mislocalization or unequal inheritance, which could have profound consequences for the long-range propagation of polarity. Here, we show that when mouse epidermal basal progenitors divide PCP components are selectively internalized into endosomes, which are inherited equally by daughter cells. Following mitosis, PCP proteins are recycled to the cell surface, where asymmetry is re-established by a process reliant on neighbouring PCP. A cytoplasmic dileucine motif governs mitotic internalization of atypical cadherin Celsr1, which recruits Vang2 and Fzd6 to endosomes. Moreover, embryos transgenic for a Celsr1 that cannot mitotically internalize exhibit perturbed hair-follicle angling, a hallmark of defective PCP. This underscores the physiological relevance and importance of this mechanism for regulating polarity during cell division.     

Neuronal polarity.

The axonal and somatodendritic domains of neurons differ in their cytoskeletal and membrane composition, complement of organelles, and capacity for macromolecular synthesis. Recently there has been progress in elucidating the cellular mechanisms that underlie the establishment and maintenance of neuronal polarity, including microtubule organization and the sorting, transport, and anchoring of membrane proteins.     

Regulation of neuronal polarity

The distinctive polarized morphology of neuronal cells is essential for the proper wiring of the nervous system. The rodent hippocampal neuron culture established about three decades ago has provided an amenable in vitro system to uncover the molecular mechanisms underlying neuronal polarization, a process relying on highly regulated cytoskeletal dynamics, membrane traffic and localized protein degradation. More recent research in vivo has highlighted the importance of the extracellular environment and cell–cell interactions in neuronal polarity. Here, I will review some key signaling pathways regulating neuronal polarization and provide some insights on the complexity of this process gained from in vivo studies.