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Summary Localized clathrin mediated endocytosis is vital for secretion and wall deposition in apical growing plant cells. Adaptor and signalling proteins, along with phosphoinositides, are known to play a regulatory, yet poorly defined role in this process. Here we investigated the function of Arabidopsis ECA4 and EPSIN3, putative mediators of the process, in pollen tubes and root hairs. Homozygous eca4 and epsin3 plants exhibited altered pollen tube morphology (in vitro) and self‐pollination led to fewer seeds and shorter siliques. These effects were augmented in eca4/epsin3 double mutant and quantitative polymerase chain reaction data revealed changes in phosphoinositide metabolism and flowering genes suggestive of a synergistic action. No visible changes were observed in root morphology, but atomic force microscopy in mutant root hairs showed altered structural stiffness. Imaging and FRET‐FLIM analysis of ECA4 and EPSIN3 X‐FP constructs revealed that both proteins interact at the plasma membrane but exhibit slightly different intracellular localization. FT‐ICR‐MS metabolomic analysis of mutant cells showed changes in lipids, amino acids and carbohydrate composition consistent with a role in secretion and growth. Characterization of double mutants of eca4 and epsin3 with phospholipase C genes (plc5, plc7) indicates that phosphoinositides (e.g. PtdIns(4,5)P2) are fundamental for a combined and complementary role of ECA4‐EPSIN3 in cell secretion. [ABSTRACT FROM AUTHOR]

Keymessage: The putative myristoylome of moss P. patens opens an avenue for studying myristoylation substrates in non-canonical model plants. A myristoylation signal was shown sufficient for membrane targeting and useful for membrane dynamics visualization during cell growth. N-myristoylation (MYR) is one form of lipid modification catalyzed by N-myristoyltransferase that enables protein-membrane association. MYR is highly conserved in all eukaryotes. However, the study of MYR is limited to a few models such as yeasts, humans, and Arabidopsis. Here, using prediction tools, we report the characterization of the putative myristoylome of the moss Physcomitrium patens. We show that basal land plants display a similar signature of MYR to Arabidopsis and may have organism-specific substrates. Phylogenetically, MYR signals have mostly co-evolved with protein function but also exhibit variability in an organism-specific manner. We also demonstrate that the MYR motif of a moss brassinosteroid-signaling kinase is an efficient plasma membrane targeting signal and labels lipid-rich domains in tip-growing cells. Our results provide insights into the myristoylome in a basal land plant and lay the foundation for future studies on MYR and its roles in plant evolution. [ABSTRACT FROM AUTHOR]

A NIMA-related protein kinase, MpNEK1, directs tip growth of rhizoids through microtubule depolymerization in a liverwort Marchantia polymorpha. The Mpnek1 knockouts were shown to develop curly and spiral rhizoids due to the fluctuated direction of growth. Still, physiological roles and mechanisms of MpNEK1-dependent rhizoid tip growth remain to be clarified. Here, we developed novel culture methods to further study rhizoid growth of M. polymorpha, in which plants were grown on vertical plates. We applied the established methods to investigate MpNEK1 function in rhizoid growth. Rhizoids of the wild-type and Mpnek1 plants grew toward the gravity. The aerial rhizoids were longer in Mpnek1 than in the wild type. When the rhizoids were grown on the surface of a cellophane sheet, rhizoid length was comparable between the wild type and Mpnek1, whereas Mpnek1 developed more rhizoids compared to the wild type. We also applied gellan gum, which is more transparent than agar, to analyze rhizoids grown in the medium. Rhizoids of Mpnek1 displayed defect on entering into the solid medium. These results suggest that Mpnek1 rhizoids have the deficiency in invasive tip growth. Thus, stable directional growth is important for rhizoids to get into the soil to anchor plant body and to adsorb water and nutrients. Collectively, our newly designed growth systems are valuable for analyzing rhizoid growth.

Introduction: VPS45 belongs to the Sec1/Munc18 family of proteins, which interact with and regulate Qa-SNARE function during membrane fusion. We have shown previously that Arabidopsis thaliana VPS45 interacts with the SYP61/SYP41/VTI12 SNARE complex, which locates on the trans-Golgi network (TGN). It is required for SYP41 stability, and it functions in cargo trafficking to the vacuole and in cell expansion. It is also required for correct auxin distribution during gravitropism and lateral root growth. Results: As vps45 knockout mutation is lethal in Arabidopsis, we identified a mutant, vps45-3, with a point mutation in the VPS45 gene causing a serine 284- to-phenylalanine substitution. The VPS45-3 protein is stable and maintains interaction with SYP61 and SYP41. However, vps45-3 plants display severe growth defects with significantly reduced organ and cell size, similar to vps45 RNAi transgenic lines that have reduced VPS45 protein levels. Root hair and pollen tube elongation, both processes of tip growth, are highly compromised in vps45-3. Mutant root hairs are shorter and thicker than those of wild-type plants, and are wavy. These root hairs have vacuolar defects, containing many small vacuoles, compared with WT root hairs with a single large vacuole occupying much of the cell volume. Pollen tubes were also significantly shorter in vps45-3 compared to WT. Discussion: We thus show that VPS45 is essential for proper tip growth and propose that the observed vacuolar defects lead to loss of the turgor pressure needed for tip growth. [ABSTRACT FROM AUTHOR]

Brown algae are complex multicellular eukaryotes whose cells possess a cell wall, which is an important structure that regulates cell size and shape. Alginate and fucose-containing sulfated polysaccharides (FCSPs) are two carbohydrate types that have major roles in influencing the mechanical properties of the cell wall (i.e. increasing or decreasing wall stiffness), which in turn regulate cell expansion, division, adhesion, and other processes; however, how brown algal cell wall structure regulates its mechanical properties, and how this relationship influences cellular growth and organismal development, is not well-understood. This chapter is focused on reviewing what we currently know about how the roles of alginates and FCSPs in brown algal developmental processes, as well as how they influence the structural and mechanical properties of cell walls. Additionally, we discuss how brown algal mutants may be leveraged to learn more about the underlying mechanisms that regulate cell wall structure, mechanics, and developmental processes, and finally we propose questions to guide future research with the use of emerging technologies. [ABSTRACT FROM AUTHOR]

Soft systems that respond to external stimuli, such as heat, magnetic field, and light, find applications in a range of fields including soft robotics, energy harvesting, and biomedicine. However, most of the existing systems exhibit nondirectional, nastic movement as they can neither grow nor sense the direction of stimuli. In this regard, artificial systems are outperformed by organisms capable of directional growth in response to the sense of stimuli or tropic growth. Inspired by tropic growth schemes of plant cells and fungal hyphae, here we report an artificial multistimuli-responsive tropic tip-growing system based on nonsolvent-induced phase separation of polymer solution, where polymer precipitates as its solvent dissolves into surrounding nonsolvent. We provide a theoretical framework to predict the size and velocity of growing precipitates and demonstrate its capability of sensing the directions of gravity, mechanical contact, and light and adjusting its growing direction in response. Exploiting the embedded physical intelligence of sensing and responding to external stimuli, our soft material system achieves multiple tasks including printing 3D structures in a confined space, bypassing mechanical obstacles, and shielded transport of liquids within water. [ABSTRACT FROM AUTHOR]

A NIMA-related protein kinase, MpNEK1, directs tip growth of rhizoids through microtubule depolymerization in a liverwort Marchantia polymorpha. The Mpnek1 knockouts were shown to develop curly and spiral rhizoids due to the fluctuated direction of growth. Still, physiological roles and mechanisms of MpNEK1-dependent rhizoid tip growth remain to be clarified. Here, we developed novel culture methods to further study rhizoid growth of M. polymorpha, in which plants were grown on vertical plates. We applied the established methods to investigate MpNEK1 function in rhizoid growth. Rhizoids of the wild-type and Mpnek1 plants grew toward the gravity. The aerial rhizoids were longer in Mpnek1 than in the wild type. When the rhizoids were grown on the surface of a cellophane sheet, rhizoid length was comparable between the wild type and Mpnek1, whereas Mpnek1 developed more rhizoids compared to the wild type. We also applied gellan gum, which is more transparent than agar, to analyze rhizoids grown in the medium. Rhizoids of Mpnek1 displayed defect on entering into the solid medium. These results suggest that Mpnek1 rhizoids have the deficiency in invasive tip growth. Thus, stable directional growth is important for rhizoids to get into the soil to anchor plant body and to adsorb water and nutrients. Collectively, our newly designed growth systems are valuable for analyzing rhizoid growth. [ABSTRACT FROM AUTHOR]

IntroductionVPS45 belongs to the Sec1/Munc18 family of proteins, which interact with and regulate Qa-SNARE function during membrane fusion. We have shown previously that Arabidopsis thaliana VPS45 interacts with the SYP61/SYP41/VTI12 SNARE complex, which locates on the trans-Golgi network (TGN). It is required for SYP41 stability, and it functions in cargo trafficking to the vacuole and in cell expansion. It is also required for correct auxin distribution during gravitropism and lateral root growth.ResultsAs vps45 knockout mutation is lethal in Arabidopsis, we identified a mutant, vps45-3, with a point mutation in the VPS45 gene causing a serine 284-to-phenylalanine substitution. The VPS45-3 protein is stable and maintains interaction with SYP61 and SYP41. However, vps45-3 plants display severe growth defects with significantly reduced organ and cell size, similar to vps45 RNAi transgenic lines that have reduced VPS45 protein levels. Root hair and pollen tube elongation, both processes of tip growth, are highly compromised in vps45-3. Mutant root hairs are shorter and thicker than those of wild-type plants, and are wavy. These root hairs have vacuolar defects, containing many small vacuoles, compared with WT root hairs with a single large vacuole occupying much of the cell volume. Pollen tubes were also significantly shorter in vps45-3 compared to WT.DiscussionWe thus show that VPS45 is essential for proper tip growth and propose that the observed vacuolar defects lead to loss of the turgor pressure needed for tip growth.

Pollen germination and subsequent pollen tube elongation are essential for successful land plant reproduction. These processes are achieved through well-documented activation of membrane trafficking and cell metabolism. Despite this, our knowledge of the dynamics of cellular phospholipids remains scarce. Here we present the turnover of the glycerolipid composition during the establishment of cell polarity and elongation processes in tobacco pollen and show the lipid composition of pollen plasma membrane-enriched fraction for the first time. To achieve this, we have combined several techniques, such as lipidomics, plasma membrane isolation, and live-cell microscopy, and performed a study with different time points during the pollen germination and pollen tube growth. Our results showed that tobacco pollen tubes undergo substantial changes in their whole-cell lipid composition during the pollen germination and growth, finding differences in most of the glycerolipids analyzed. Notably, while lysophospholipid levels decrease during germination and growth, phosphatidic acid increases significantly at cell polarity establishment and continues with similar abundance in cell elongation. We corroborated these findings by measuring several phospholipase activities in situ. We also observed that lysophospholipids and phosphatidic acid are more abundant in the plasma membrane-enriched fraction than that in the whole cell. Our results support the important role for the phosphatidic acid in the establishment and maintenance of cellular polarity in tobacco pollen tubes and indicate that plasma membrane lysophospholipids may be involved in pollen germination. [ABSTRACT FROM AUTHOR]

Summary: Polar cell growth in plants requires a cell peripheral region that undergoes membrane extension and cell wall remodeling. Since the 1990s, RHO‐RELATED GTPASES FROM PLANTS (ROPs) have been identified as master regulators that determine the site of cell growth. ROPs function to regulate actin and microtubule cytoskeletons, calcium gradients, and exocytosis, thus directing the delivery of materials for membrane and cell wall extension. In recent years, our understanding of the regulatory mechanisms underlying polar localization and the activation of ROPs has greatly advanced. Evidence points to the crucial roles of membrane lipids, receptor‐like kinases, and cell wall components. In this review, we provide updates on the mechanisms underlying polarity control in tip‐growing cells, with a focus on ROP effectors and membrane‐associated signals. By integrating knowledge from pollen tubes, root hairs, and findings in bryophyte protonema cells and rhizoids, we hope to offer important insights into a common conceptual framework on polarity establishment governed by intercellular and extracellular signals. [ABSTRACT FROM AUTHOR]

The red alga Neopyropia yezoensis undergoes polarized elongation and asymmetrical cell division of the apical stem cell during tip growth in filamentous generations of its life cycle: the conchocelis and conchosporangium. Side branches are also produced via tip growth, a process involving the regeneration and asymmetrical division of the apical stem cell. Here, we demonstrate that auxin plays a crucial role in these processes by using the auxin antagonist 2-(1H-Indol-3-yl)-4-oxo-4-phenyl-butyric acid (PEO-IAA), which specifically blocks the activity of the auxin receptor TRANSPORT INHIBITOR RESPONSE1 (TIR1) in land plants. PEO-IAA repressed both the regeneration and polarized tip growth of the apical stem cell in single-celled conchocelis; this phenomenon was reversed by treatment with the auxin indole-3-acetic acid (IAA). In addition, tip growth of the conchosporangium was accelerated by IAA treatment but repressed by PEO-IAA treatment. These findings indicate that auxin regulates polarized tip cell growth and that an auxin receptor-like protein is present in N. yezoensis. The sensitivity to different 5-alkoxy-IAA analogs differs considerably between N. yezoensis and Arabidopsis thaliana. N. yezoensis lacks a gene encoding TIR1, indicating that its auxin receptor-like protein differs from the auxin receptor of terrestrial plants. These findings shed light on auxin-induced mechanisms and the regulation of tip growth in plants. [ABSTRACT FROM AUTHOR]

Deep mycosis is a severe fungal disease that could result in fatal outcomes. However, there is still a demand for highly effective and safe antifungal drugs, given the side effects of the existing treatments and the increase in the resistance to them. In this study, we evaluated the involvement of the lipid kinase Fab1 and its activator Vac14 (Fab1/Vac14) in tip growth in Saccharomyces cerevisiae INVSc1, along with their impact on cell proliferation, using a genetic approach to inhibit them. The results revealed that Fab1/Vac14 inhibition suppressed growth and caused an increase in the rate of β-1,3-glucanase (BGL2) fused with emerald green fluorescent protein (EmGFP) (BGL2-EmGFP) localization at the tip. The inhibition of the endocytic pathway using a lysosome inhibitor also resulted in an increased localization of BGL2-EmGFP at the tip. The overexpression of wild-type BGL2-EmGFP, but not that of the inactive mutant BGL2, led to a complete loss of the cell proliferation ability. These findings suggested that the Fab1/Vac14 complex could be a novel target for the development of antifungal drugs based on tip growth regulation, possibly via excessive cell wall degradation., Competing Interests: Declaration of competing interest The authors declare the following financial interests/personal relationships which may be considered as potential competing interests: Sen Takeshita reports a relationship with Ajinomoto Co., Inc. that includes: employment. If there are other authors, they declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024. Published by Elsevier Inc.)

A central problem of geometry is the tiling of space with simple structures. The classical solutions, such as triangles, squares, and hexagons in the plane and cubes and other polyhedra in three-dimensional space are built with sharp corners and flat faces. However, many tilings in Nature are characterized by shapes with curved edges, nonflat faces, and few, if any, sharp corners. An important question is then to relate prototypical sharp tilings to softer natural shapes. Here, we solve this problem by introducing a new class of shapes, the soft cells , minimizing the number of sharp corners and filling space as soft tilings . We prove that an infinite class of polyhedral tilings can be smoothly deformed into soft tilings and we construct the soft versions of all Dirichlet-Voronoi cells associated with point lattices in two and three dimensions. Remarkably, these ideal soft shapes, born out of geometry, are found abundantly in nature, from cells to shells., (© The Author(s) 2024. Published by Oxford University Press on behalf of National Academy of Sciences.)

Root hairs are tubular outgrowths of epidermal cells that increase the root surface area and thereby make the root more efficient at absorbing water and nutrients. Their expansion is limited to the root hair apex, where growth is reported to take place in a pulsating manner. These growth pulses coincide with oscillations of the apoplastic and cytosolic pH in a similar way as has been reported for pollen tubes. Likewise, the concentrations of apoplastic reactive oxygen species (ROS) and cytoplasmic Ca2+ oscillate with the same periodicity as growth. Whereas ROS appear to control cell wall extensibility and opening of Ca2+ channels, the role of protons as a growth signal in root hairs is less clear andmay differ fromthat in pollen tubes where plasmamembrane H+-ATPases have been shown to sustain growth. In this review, we outline our current understanding of how pH contributes to root hair development. [ABSTRACT FROM AUTHOR]

In plant cells, linkage between the cytoskeleton, plasma membrane, and cell wall is crucial for maintaining cell shape. In highly polarized pollen tubes, this coordination is especially important to allow rapid tip growth and successful fertilization. Class I formins contain cytoplasmic actin-nucleating formin homology domains as well as a proline-rich extracellular domain and are candidate coordination factors. Here, using Arabidopsis, we investigated the functional significance of the extracellular domain of two pollen-expressed class I formins: AtFH3, which does not have a polar localization, and AtFH5, which is limited to the growing tip region. We show that the extracellular domain of both is necessary for their function, and identify distinct O -glycans attached to these sequences, AtFH5 being hydroxyproline-arabinosylated and AtFH3 carrying arabinogalactan chains. Loss of hydroxyproline arabinosylation altered the plasma membrane localization of AtFH5 and disrupted actin cytoskeleton organization. Moreover, we show that O -glycans differentially affect lateral mobility in the plasma membrane. Together, our results support a model of protein sub-functionalization in which AtFH5 and AtFH3, restricted to specific plasma membrane domains by their extracellular domains and the glycans attached to them, organize distinct subarrays of actin during pollen tube elongation. [ABSTRACT FROM AUTHOR]

Tip growth is an extreme form of polarized cell expansion that occurs in all eukaryotic kingdoms to generate highly elongated tubular cells with specialized functions, including fungal hyphae, animal neurons, plant pollen tubes, and root hairs (RHs). RHs are tubular structures that protrude from the root epidermis to facilitate water and nutrient uptake, microbial interactions, and plant anchorage. RH tip growth requires polarized vesicle targeting and active exocytosis at apical growth sites. However, how apical exocytosis is spatially and temporally controlled during tip growth remains elusive. Here, we report that the Qa-Soluble N -ethylmaleimide-sensitive factor attachment protein receptor (SNARE) SYP121 acts as an effector of Rho of Plants 2 (ROP2), mediating the regulation of RH tip growth. We show that active ROP2 promotes SYP121 targeting to the apical plasma membrane. Moreover, ROP2 directly interacts with SYP121 and promotes the interaction between SYP121 and the R-SNARE VAMP722 to form a SNARE complex, probably by facilitating the release of the Sec1/Munc18 protein SEC11, which suppresses the function of SYP121. Thus, the ROP2-SYP121 pathway facilitates exocytic trafficking during RH tip growth. Our study uncovers a direct link between an ROP GTPase and vesicular trafficking and a new mechanism for the control of apical exocytosis, whereby ROP GTPase signaling spatially regulates SNARE complex assembly and the polar distribution of a Q-SNARE. Root hair tip growth requires polarized vesicle targeting and active exocytosis at apical growth site, but how it is spatially and temporally controlled remains elusive. This study demonstrates that ROP2 directly regulates the polar localization of SYP121 and its interaction with VAMP722 for SNARE complex assembly, establishing a tight link between ROP signaling and apical exocytosis during root hair tip growth. [ABSTRACT FROM AUTHOR]

Key message: The pollen tube is being subjected to control by a complex network of communication that regulates its shape and the misfunction of a single component causes specific deformations. In flowering plants, the pollen tube is a tubular extension of the pollen grain required for successful sexual reproduction. Indeed, maintaining the unique shape of the pollen tube is essential for the pollen tube to approach the embryo sac. Many processes and molecules (such as GTPase activity, phosphoinositides, Ca2+ gradient, distribution of reactive oxygen species and nitric oxide, nonuniform pH values, organization of the cytoskeleton, balance between exocytosis and endocytosis, and cell wall structure) play key and coordinated roles in maintaining the cylindrical shape of pollen tubes. In addition, the above factors must also interact with each other so that the cell shape is maintained while the pollen tube follows chemical signals in the pistil that guide it to the embryo sac. Any intrinsic changes (such as erroneous signals) or extrinsic changes (such as environmental stresses) can affect the above factors and thus fertilization by altering the tube morphology. In this review, the processes and molecules that enable the development and maintenance of the unique shape of pollen tubes in angiosperms are presented emphasizing their interaction with specific tube shape. Thus, the purpose of the review is to investigate whether specific deformations in pollen tubes can help us to better understand the mechanism underlying pollen tube shape. [ABSTRACT FROM AUTHOR]

Root hairs are tubular outgrowths of epidermal cells that increase the root surface area and thereby make the root more efficient at absorbing water and nutrients. Their expansion is limited to the root hair apex, where growth is reported to take place in a pulsating manner. These growth pulses coincide with oscillations of the apoplastic and cytosolic pH in a similar way as has been reported for pollen tubes. Likewise, the concentrations of apoplastic reactive oxygen species (ROS) and cytoplasmic Ca2+ oscillate with the same periodicity as growth. Whereas ROS appear to control cell wall extensibility and opening of Ca2+ channels, the role of protons as a growth signal in root hairs is less clear and may differ from that in pollen tubes where plasma membrane H+-ATPases have been shown to sustain growth. In this review, we outline our current understanding of how pH contributes to root hair development.

The use of Ni-based catalysts is a common method for eliminating tar through catalytic cracking. Carbon deposition is the main cause of deactivation in Ni/ZSM-5 catalysts, with filamentous MWCNTs being the primary form of carbon deposits. This study investigates the formation and evolution of CNTs during the catalytic process of biomass tar to explore the mechanism behind carbon deposition. The effect of the 9Ni/10MWCNTs/81ZSM-5 on toluene reforming was investigated through a vertical furnace. Gases produced by tar catalysis were evaluated through GC analysis. The physicochemical structure, properties and catalytic performance of the catalyst were also tested. TG analysis was used to assess the accumulation and oxidation reactivity of carbon on the catalyst surface. An analysis was conducted on the mechanism of carbon deposition during catalyst deactivation in tar catalysis. The results showed that the 9Ni/91ZSM-5 had a superior toluene conversion of 60.49%, but also experienced rapid and substantial carbon deposition up to 52.69%. Carbon is mainly deposited as curved filaments on both the surface and pore channels of the catalyst. In some cases, tip growth occurs where both carbon deposition and Ni coexist. Furthermore, specific surface area and micropore volume are reduced to varying degrees due to carbon deposition. With the time increased, the amount of carbon deposited on the catalyst surface increased to 62.81%, which gradually approached saturation, and the overall performance of the catalyst was stabilized. This situation causes toluene molecules to detach from the active sites within the catalyst, hindering gas release, which leads to reduced catalytic activity and further carbon deposition. It provides both a basis for the development of new catalysts and an economically feasible solution for practical tar reduction and removal. [Display omitted] • The carbonaceous deposition during 9Ni/91ZSM-5 catalyzed toluene were studied. • The effect of carbon nanotube content on toluene conversion were analyzed. • Investigating the surface morphology and structure of catalyst after catalysis. • Carbon deposition and oxidative reactivity were explored for effecting of catalyst. • Proposed generation mechanism of carbon during toluene catalyzed by 9Ni/91ZSM-5. [ABSTRACT FROM AUTHOR]

In this study, a growth mechanism of vertically aligned carbon nanotubes (VACNT) on a nanostructured porous silicon template (NPSiT) was proposed based on the experimental observations obtained. Camphor oil and ferrocene (Fe) were used as a carbon source and a catalyst, respectively. The NPSiT was optimised and used as a template to synthesise VACNT. The VACNT was synthesised on NPSiT by using a double thermal chemical vapour deposition (DTCVD) method. The synthesis was also performed via a floated method in a DTCVD reactor by using an in-situ catalyst in the absence of any catalysts pre-treatment. Each of the growth mechanisms was proposed based on the TEM images obtained at different synthesis conditions. From the results, a mixed top and bottom growth model was proposed for the growth mechanism of VACNT at the pores and pillars. On the contrary, the tip growth model was proposed for the growth inside the pores, while the valley growth model was proposed for the growth within the pillars of NPSiT. Furthermore, a mixed tip and valley growth model was proposed based on the location of Fe particles. [ABSTRACT FROM AUTHOR]

Summary: Small signalling peptides are key molecules for cell‐to‐cell communications in plants. The cysteine‐rich signalling peptide, rapid alkalinisation factors (RALFs) family are involved in diverse developmental and stress responses and have expanded considerably during land plant evolution, implying neofunctionalisations in the RALF family. However, the ancestral roles of RALFs when land plant first acquired them remain unknown.Here, we functionally characterised two of the three RALFs in bryophyte Physcomitrium patens using loss‐of‐function mutants, overexpressors, as well as fluorescent proteins tagged reporter lines.We showed that PpRALF1 and PpRALF2 have overlapping functions in promoting protonema tip growth and elongation, showing a homologous function as the Arabidopsis RALF1 in promoting root hair tip growth. Although both PpRALFs are secreted to the plasma membrane on which PpRALF1 symmetrically localised, PpRALF2 showed a polarised localisation at the growing tip. Notably, proteolytic cleavage of PpRALF1 is necessary for its function.Our data reveal a possible evolutionary origin of the RALF functions and suggest that functional divergence of RALFs is essential to drive complex morphogenesis and to facilitate other novel processes in land plants. [ABSTRACT FROM AUTHOR]

Root hairs are cylindrical extensions of root epidermal cells that are important for the acquisition of water and minerals, interactions between plant and microbes. The deposition of cell wall materials in the tip enables root hairs to maintain elongation constantly. To date, our knowledge of the regulators that connect the architecture of cell wall and the root hair development remains very limited. Here, we demonstrated that COBL9 and COBL7 , two genes of COBRA-Like family in Arabidopsis as well as their counterparts in rice, OsBC1L1 and OsBC1L8, regulate root hair growth. Single mutant cobl9 , double mutants cobl7 cobl9 and double mutants osbc1l1 osbc1l8 all displayed prematurely terminated root hair elongation, though at varying levels. COBL7-YFP and COBL9-YFP accumulate prominently in the growing tips of newly emerged root hairs. Furthermore, cobl9, cobl7 cobl9 and osbc1l1 osbc1l8 mutants were defective in the enrichment of cellulose in the tips of the growing root hairs. We also discovered that overexpression of COBL9 could promote root hair elongation and salinity tolerance. Taken together, these results provide compelling evidence that the polarized COBL7 and COBL9 in the tip of the emerging root hairs have conserved roles in regulating root hair development and stress adaptation in dicots and monocots. • COBL9 and COBL7 synergistically regulate root hair tip growth through controlling apical cellulose deposition in Arabidopsis. • COBL7 and COBL9 have conserved roles in regulating root hair tip growth between dicots and monocots. [ABSTRACT FROM AUTHOR]

In the process of pollination, a pollen tube grows from a pollen grain that has fallen on the stigma of a flower. This tube grows towards the ovary of the flower where it will deliver male reproductive material. Knowledge of the dynamics of pollen tube growth will provide a basis for understanding more complex cells that exhibit similar growth behavior. Current pollen tube growth models are a collection of differential equations that represent the level of understanding that biologists have concerning apical growth. Due to their complex nature, these models are not used to verify observed behavior in living cells as seen under a microscope. We present a model that can be used to describe the behavior of growing pollen tube cells in actual experiments. We propose biologically relevant functions based on knowledge of the growth process to explain the dynamics of model parameters. Our model uses an affine transformation to propagate the tip of the cell and statistical parameter estimation to measure necessary parameters during growth. Using experimental videos of pollen tube growth, we show that our model can adapt to various growth scenarios while extracting growth parameters from the videos.

In the process of pollination, a pollen tube grows from a pollen grain that has fallen on the stigma of a flower. This tube grows towards the ovary of the flower where it will deliver male reproductive material. Knowledge of the dynamics of pollen tube growth will provide a basis for understanding more complex cells that exhibit similar growth behavior. Current pollen tube growth models are a collection of differential equations that represent the level of understanding that biologists have concerning apical growth. Due to their complex nature, these models are not used to verify observed behavior in living cells as seen under a microscope. We present a model that can be used to describe the behavior of growing pollen tube cells in actual experiments. We propose biologically relevant functions based on knowledge of the growth process to explain the dynamics of model parameters. Our model uses an affine transformation to propagate the tip of the cell and statistical parameter estimation to measure necessary parameters during growth. Using experimental videos of pollen tube growth, we show that our model can adapt to various growth scenarios while extracting growth parameters from the videos.

A wide range of tip-growing cells in plants display polarized cell growth, which is an essential cellular process for the form and function of individual cells. Understanding of the regulatory mechanisms underlying tip growth in terrestrial plants has improved. Cellular processes involved in tip growth have also been investigated in some algae species that form filamentous cells, but their regulatory mechanisms remain unclear. In the macro red alga Neopyropia yezoensis , for which genome information has recently been released, the conchocelis apical cell exhibits tip growth and forms a filamentous structure. Here, we report a live-imaging technique using high-resolution microscopy to analyze the tip growth and cell division of N. yezoensis conchocelis. This imaging analysis addressed tip growth dynamics and cell division in conchocelis apical cells. The directionality and tip growth expansion were disrupted by the application of cytoskeletal drugs, suggesting the involvement of microtubules (MTs) and actin filaments (AFs) in these processes. A growing apical cell mostly contained a single chloroplast that moved toward the expanding part of the apical cell. Drug application also inhibited chloroplast movement, implying that the movement may be dependent on the cytoskeleton. The study determined that live-imaging analysis is a versatile approach for exploring the dynamics of tip growth and cell division in N. yezoensis conchocelis, which provides insights into the regulatory mechanisms underlying cellular growth in multicellular red algae. [ABSTRACT FROM AUTHOR]

Pollen grains transport the sperm cells through the style tissue via a fast-growing pollen tube to the ovaries where fertilization takes place. Pollen tube growth requires a precisely regulated network of cellular as well as molecular events including the activity of the plasma membrane H+ ATPase, which is known to be regulated by reversible protein phosphorylation and subsequent binding of 14-3-3 isoforms. Immunodetection of the phosphorylated penultimate threonine residue of the pollen plasma membrane H+ ATPase (LilHA1) of Lilium longiflorum pollen revealed a sudden increase in phosphorylation with the start of pollen tube growth. In addition to phosphorylation, pH modulated the binding of 14-3-3 isoforms to the regulatory domain of the H+ ATPase, whereas metabolic components had only small effects on 14-3-3 binding, as tested with in vitro assays using recombinant 14-3-3 isoforms and phosphomimicking substitutions of the threonine residue. Consequently, local H+ influxes and effluxes as well as pH gradients in the pollen tube tip are generated by localized regulation of the H+ ATPase activity rather than by heterogeneous localized distribution in the plasma membrane. [ABSTRACT FROM AUTHOR]

A wide range of tip-growing cells in plants display polarized cell growth, which is an essential cellular process for the form and function of individual cells. Understanding of the regulatory mechanisms underlying tip growth in terrestrial plants has improved. Cellular processes involved in tip growth have also been investigated in some algae species that form filamentous cells, but their regulatory mechanisms remain unclear. In the macro red alga Neopyropia yezoensis, for which genome information has recently been released, the conchocelis apical cell exhibits tip growth and forms a filamentous structure. Here, we report a live-imaging technique using high-resolution microscopy to analyze the tip growth and cell division of N. yezoensis conchocelis. This imaging analysis addressed tip growth dynamics and cell division in conchocelis apical cells. The directionality and tip growth expansion were disrupted by the application of cytoskeletal drugs, suggesting the involvement of microtubules (MTs) and actin filaments (AFs) in these processes. A growing apical cell mostly contained a single chloroplast that moved toward the expanding part of the apical cell. Drug application also inhibited chloroplast movement, implying that the movement may be dependent on the cytoskeleton. The study determined that live-imaging analysis is a versatile approach for exploring the dynamics of tip growth and cell division in N. yezoensis conchocelis, which provides insights into the regulatory mechanisms underlying cellular growth in multicellular red algae.

In this protocol, we focused on analyzing internal branches of Drosophila class IV neurons. These neurons are characterized by their highly branched axons and dendrites and intricately tile the larval body. As Drosophila larvae progress through developmental stages, the dendritic arbors of Class IV neurons undergo notable transformations. As Drosophila larvae develop, their Class IV dendritic arbors grow. In the initial 24 h after egg laying (AEL), the dendrites are smaller than segments. During the subsequent 24 h of the first instar larval stage, dendritic arbors outpace segment growth, achieving tiling. After 48 h, arbors and segments grow concurrently. Epidermal cells near Class IV dendrites expand in proportion to segment growth. This observation suggested that Class IV cells might grow via branch dilation-uniformly elongating branches, akin to Class I cells [1,2]. To understand whether the class IV complex arbor structure is formed by dilation or simply from growing tips, we developed this protocol to introduce a systematic approach for quantitatively assessing the growth dynamics of internal branches. Key features • This protocol employs imaging the same neuron over different development times • Drosophila embryo and larvae genotype is ;;ppkCD4-tdGFP, which explicitly tags class IV neurons • This protocol for the preparation of agar pads to mount and image Drosophila larvae is adapted from Monica Driscoll's method • Neurons are imaged without the use of anesthetics and for a short duration of time • This technique involves the use of a spinning disk confocal microscope., Competing Interests: Competing interestsThe authors declare that they have no competing interests., (©Copyright : © 2024 The Authors; This is an open access article under the CC BY-NC license.)

Hydrogel tubes made of sodium alginate (SA) have potential applications in drug delivery, soft robots, biomimetic blood vessels, tissue stents, and other fields. However, the continuous preparation of hollow SA hydrogel tubes with good stability and size control remains a huge challenge for chemists, material scientists, and medical practitioners. Inspired by the plant apical growth strategy, a new method named soft cap-guided growth was proposed to produce SA hydrogel tubes. Due to the introduction of inert low gravity substances, such as air and heptane, into the extrusion needle in front of calcium chloride solution to form a soft cap, the SA hydrogel tubes with controllable sizes were fabricated rapidly and continuously without using a template through a negative gravitropism mechanism. The SA hydrogel tubes had good tensile strength, high burst pressure, and good cell compatibility. In addition, hydrogel tubes with complex patterns were conveniently created by controlling the motion path of a soft cap, such as a rotating SA bath or magnetic force. Our research provided a simple innovative technique to steer the growth of hydrogel tubes, which made it possible to mass produce hydrogel tubes with controllable sizes and programmable patterns.

Summary: All kingdoms of life have evolved tip‐growing cells able to mine their environment or deliver cargo to remote targets. The basic cellular processes supporting these functions are understood in increasing detail, but the multiple interactions between them lead to complex responses that require quantitative models to be disentangled. Here, I review the equations that capture the fundamental interactions between wall mechanics and cell hydraulics starting with a detailed presentation of James Lockhart's seminal model. The homeostatic feedbacks needed to maintain a steady tip velocity are then shown to offer a credible explanation for the pulsatile growth observed in some tip‐growing cells. Turgor pressure emerges as a central variable whose role in the morphogenetic process has been a source of controversy for more than 50 yr. I argue that recasting Lockhart's work as a process of chemical stress relaxation can clarify how cells control tip growth and help us internalise the important but passive role played by turgor pressure in the morphogenetic process. [ABSTRACT FROM AUTHOR]

Polydimethylsiloxane (PDMS) chips have proven to be suitable environments for the growth of several filamentous organisms. However, depending on the specimen, the number of investigations concerning their growth and cell differentiation is limited. In this work, we monitored the developmental pattern of the brown alga Ectocarpus inside PDMS lab-on-chips. Two main methods of inoculation of the lab-on-chip were tested, i.e., via the direct injection of spores into the chamber as well as through the insertion of sporophyte filaments. The resulting growth rate, growth trajectory, cell differentiation, and cell branching were monitored and quantified for 20 days inside 25 or 40 μm parallel channels under standard light and temperature conditions. With growth rates of 2.8 μm⋅h–1, normal growth trajectories and cell differentiation, as well as branching occurring inside the microfluidic environment, the main development steps were shown to be similar to those observed in non-constrained in vitro conditions. Additionally, the labelling of Ectocarpus cell wall polysaccharides using calcofluor for cellulose detection and immunolocalisation with monoclonal antibodies for alginates showed the expected patterns when compared to open space growth evaluated with either epifluorescence or confocal microscopy. Overall, this article describes the experimental conditions for observing and studying the basic unaltered processes of brown algal growth using microfluidic technology which provides the basis for future biochemical and biological researches.

Small GTPase is a type of crucial regulator in eukaryotes. It acts as a molecular switch by binding with GTP and GDP in cytoplasm, affecting various cellular processes. Small GTPase were divided into five subfamilies based on sequence, structure and function: Ras, Rho, Rab, Arf/Sar and Ran, with Rab being the largest subfamily. Members of the Rab subfamily play an important role in regulating complex vesicle transport and microtubule system activity. Plant cells are composed of various membrane-bound organelles, and vesicle trafficking is fundamental to the existence of plants. At present, the function of some Rab members, such as RabA1a, RabD2b/c and RabF2, has been well characterized in plants. This review summarizes the role of Rab GTPase in regulating plant tip growth, morphogenesis, fruit ripening and stress response, and briefly describes the regulatory mechanisms involved. It provides a reference for further alleviating environmental stress, improving plant resistance and even improving fruit quality. [ABSTRACT FROM AUTHOR]

The red alga Neopyropia yezoensis undergoes polarized elongation and asymmetrical cell division of the apical stem cell during tip growth in filamentous generations of its life cycle: the conchocelis and conchosporangium. Side branches are also produced via tip growth, a process involving the regeneration and asymmetrical division of the apical stem cell. Here, we demonstrate that auxin plays a crucial role in these processes by using the auxin antagonist 2-(1H-Indol-3-yl)-4-oxo-4-phenyl-butyric acid (PEO-IAA), which specifically blocks the activity of the auxin receptor TRANSPORT INHIBITOR RESPONSE1 (TIR1) in land plants. PEO-IAA repressed both the regeneration and polarized tip growth of the apical stem cell in single-celled conchocelis; this phenomenon was reversed by treatment with the auxin indole-3-acetic acid (IAA). In addition, tip growth of the conchosporangium was accelerated by IAA treatment but repressed by PEO-IAA treatment. These findings indicate that auxin regulates polarized tip cell growth and that an auxin receptor-like protein is present in N. yezoensis. The sensitivity to different 5-alkoxy-IAA analogs differs considerably between N. yezoensis and Arabidopsis thaliana. N. yezoensis lacks a gene encoding TIR1, indicating that its auxin receptor-like protein differs from the auxin receptor of terrestrial plants. These findings shed light on auxin-induced mechanisms and the regulation of tip growth in plants.

Formins are a large, evolutionarily conserved family of actin-nucleating proteins with additional roles in regulating microfilament, microtubule, and membrane dynamics. Angiosperm formins, expressed in both sporophytic and gametophytic tissues, can be divided into two subfamilies, Class I and Class II, each often exhibiting characteristic domain organization. Gametophytically expressed Class I formins have been documented to mediate plasma membrane-based actin assembly in pollen grains and pollen tubes, contributing to proper pollen germination and pollen tube tip growth, and a rice Class II formin, FH5/RMD, has been proposed to act as a positive regulator of pollen tube growth based on mutant phenotype and overexpression data. Here we report functional characterization of the Arabidopsis thaliana pollen-expressed typical Class II formin FH13 (At5g58160). Consistent with published transcriptome data, live-cell imaging in transgenic plants expressing fluorescent protein-tagged FH13 under the control of the FH13 promoter revealed expression in pollen and pollen tubes with non-homogeneous signal distribution in pollen tube cytoplasm, suggesting that this formin functions in the male gametophyte. Surprisingly, fh13 loss of function mutations do not affect plant fertility but result in stimulation of in vitro pollen tube growth, while tagged FH13 overexpression inhibits pollen tube elongation. Pollen tubes of mutants expressing a fluorescent actin marker exhibited possible minor alterations of actin organization. Our results thus indicate that FH13 controls or limits pollen tube growth, or, more generally, that typical Class II formins should be understood as modulators of pollen tube elongation rather than merely components of the molecular apparatus executing tip growth.

Formins are a large, evolutionarily conserved family of actin-nucleating proteins with additional roles in regulating microfilament, microtubule, and membrane dynamics. Angiosperm formins, expressed in both sporophytic and gametophytic tissues, can be divided into two subfamilies, Class I and Class II, each often exhibiting characteristic domain organization. Gametophytically expressed Class I formins have been documented to mediate plasma membrane-based actin assembly in pollen grains and pollen tubes, contributing to proper pollen germination and pollen tube tip growth, and a rice Class II formin, FH5/RMD, has been proposed to act as a positive regulator of pollen tube growth based on mutant phenotype and overexpression data. Here we report functional characterization of the Arabidopsis thaliana pollen-expressed typical Class II formin FH13 (At5g58160). Consistent with published transcriptome data, live-cell imaging in transgenic plants expressing fluorescent protein-tagged FH13 under the control of the FH13 promoter revealed expression in pollen and pollen tubes with non-homogeneous signal distribution in pollen tube cytoplasm, suggesting that this formin functions in the male gametophyte. Surprisingly, fh13 loss of function mutations do not affect plant fertility but result in stimulation of in vitro pollen tube growth, while tagged FH13 overexpression inhibits pollen tube elongation. Pollen tubes of mutants expressing a fluorescent actin marker exhibited possible minor alterations of actin organization. Our results thus indicate that FH13 controls or limits pollen tube growth, or, more generally, that typical Class II formins should be understood as modulators of pollen tube elongation rather than merely components of the molecular apparatus executing tip growth. [ABSTRACT FROM AUTHOR]

In eukaryotic cells, organelle and vesicle transport, positioning, and interactions play crucial roles in cytoplasmic organization and function. These processes are governed by intracellular trafficking mechanisms. At the core of that trafficking, the cytoskeleton and directional transport by motor proteins stand out as its key regulators. Plant cell tip growth is a well-studied example of cytoplasm organization by polarization. This polarization, essential for the cell's function, is driven by the cytoskeleton and its associated motors. This review will focus on myosin XI, a molecular motor critical for vesicle trafficking and polarized plant cell growth. We will center our discussion on recent data from the moss Physcomitrium patens and the liverwort Marchantia polymorpha. The biochemical properties and structure of myosin XI in various plant species are discussed, highlighting functional conservation across species. We further explore this conservation of myosin XI function in the process of vesicle transport in tip-growing cells. Existing evidence indicates that myosin XI actively organizes actin filaments in tip-growing cells by a mechanism based on vesicle clustering at their tips. A hypothetical model is presented to explain the essential function of myosin XI in polarized plant cell growth based on vesicle clustering at the tip. The review also provides insight into the in vivo localization and dynamics of myosin XI, emphasizing its role in cytosolic calcium regulation, which influences the polymerization of F-actin. Lastly, we touch upon the need for additional research to elucidate the regulation of myosin function., (© 2024 The Author(s). Published by Portland Press Limited on behalf of the Biochemical Society.)

Brown algae (Phaeophyceae) are multicellular photoautrophic organisms and the largest biomass producers in coastal regions. A variety of observations indicate that their extracellular matrix (ECM) is involved with screening of salts, development, cell fate selection, and defense responses. It is likely that these functionalities are related to its constitutive structures. The major components of the ECM of brown algae are β-glucans, alginates, and fucose-containing sulfated polysaccharides. The genus Ectocarpus comprises a wide range of species that have adapted to different environments, including isolates of Ectocarpus subulatus, a species highly resistant to low salinity. Previous studies on a freshwater strain of E. subulatus indicated that the sulfate remodeling of fucans is related to the external salt concentration. Here we show that the sulfate content of the surrounding medium is a key parameter influencing both the patterning of the alga and the occurrence of the BAM4 sulfated fucan epitope in walls of apical cells. These results indicate that sulfate uptake and incorporation in the sulfated fucans from apical cells is an essential parameter to sustain tip growth, and we discuss its influence on the architectural plasticity of Ectocarpus.

The pollen-specific calcium-dependent protein kinase PiCDPK1 of Petunia inflata has previously been shown to regulate polarity in tip growth in pollen tubes. Here we report the identification of a Rho Guanine Dissociation Inhibitor (PiRhoGDI1) as a PiCDPK1 interacting protein. We demonstrate that PiRhoGDI1 and PiCDPK1 interact in a yeast 2-hybrid assay, as well as in an in vitro pull-down assay, and that PiRhoGDI1 is phosphorylated by PiCDPK1 in vitro. We further demonstrate the PiRhoGDI1 is capable of rescuing the loss of growth polarity phenotype caused by over-expressing PiCDPK1 in vivo using stable transgenic plants. We confirmed that PiRhoGDI1 interacts with a pollen-expressed ROP GTPase isoform consistent with the established role of RhoGDIs in negatively regulating GTPases through their membrane removal and locking them in an inactive cytosolic complex. ROP is a central regulator of polarity in tip growth, upstream of Ca2+, and PiCDPK1 over-expression has been previously reported to lead to dramatic elevation of cytosolic Ca2+ through a positive feedback loop. The discovery that PiCDPK1 impacts ROP regulation via PiRhoGDI1 suggests that PiCDPK1 acts as RhoGDI displacement factor and leads us to propose a model which we hypothesize regulates the rapid recycling of ROP GTPase at the pollen tube tip.

Brown algae (Phaeophyceae) are multicellular photoautrophic organisms and the largest biomass producers in coastal regions. A variety of observations indicate that their extracellular matrix (ECM) is involved with screening of salts, development, cell fate selection, and defense responses. It is likely that these functionalities are related to its constitutive structures. The major components of the ECM of brown algae are β-glucans, alginates, and fucose-containing sulfated polysaccharides. The genus Ectocarpus comprises a wide range of species that have adapted to different environments, including isolates of Ectocarpus subulatus , a species highly resistant to low salinity. Previous studies on a freshwater strain of E. subulatus indicated that the sulfate remodeling of fucans is related to the external salt concentration. Here we show that the sulfate content of the surrounding medium is a key parameter influencing both the patterning of the alga and the occurrence of the BAM4 sulfated fucan epitope in walls of apical cells. These results indicate that sulfate uptake and incorporation in the sulfated fucans from apical cells is an essential parameter to sustain tip growth, and we discuss its influence on the architectural plasticity of Ectocarpus. [ABSTRACT FROM AUTHOR]

Summary: HYDROXYPROLINE O‐ARABINOSYLTRANSFERASEs (HPATs) initiate a post‐translational protein modification (Hyp‐Ara) found abundantly on cell wall structural proteins. In Arabidopsis thaliana, HPAT1 and HPAT3 are redundantly required for full pollen fertility. In addition to the lack of Hyp‐Ara in hpat1/3 pollen tubes (PTs), we also found broadly disrupted cell wall polymer distributions, particularly the conversion of the tip cell wall to a more shaft‐like state. Mutant PTs were slow growing and prone to rupture and morphological irregularities. In a forward mutagenesis screen for suppressors of the hpat1/3 low seed‐set phenotype, we identified a missense mutation in exo70a2, a predicted member of the vesicle‐tethering exocyst complex. The suppressed pollen had increased fertility, fewer morphological defects and partially rescued cell wall organization. A transcriptional null allele of exo70a2 also suppressed the hpat1/3 fertility phenotype, as did mutants of core exocyst complex member sec15a, indicating that reduced exocyst function bypassed the PT requirement for Hyp‐Ara. In a wild‐type background, exo70a2 reduced male transmission efficiency, lowered pollen germination frequency and slowed PT elongation. EXO70A2 also localized to the PT tip plasma membrane, consistent with a role in exocyst‐mediated secretion. To monitor the trafficking of Hyp‐Ara modified proteins, we generated an HPAT‐targeted fluorescent secretion reporter. Reporter secretion was partially dependent on EXO70A2 and was significantly increased in hpat1/3 PTs compared with the wild type, but was reduced in the suppressed exo70a2 hpat1/3 tubes. Significance Statement: Pollen tube growth requires the highly polarized secretion of new cell wall material to the pollen tube tip, but how this secreted material is incorporated into a functional cell wall remains poorly understood. Here, we demonstrate a novel functional link between protein glycosylation, cell wall organization and secretion rate. [ABSTRACT FROM AUTHOR]

Summary: Phosphatidic acid (PA), an important signalling and metabolic phospholipid, is predominantly localized in the subapical plasma membrane (PM) of growing pollen tubes. PA can be produced from structural phospholipids by phospholipase D (PLD), but the isoforms responsible for production of PM PA were not identified yet and their functional roles remain unknown. Following genome‐wide bioinformatic analysis of the PLD family in tobacco, we focused on the pollen‐overrepresented PLDδ class. Combining live‐cell imaging, gene overexpression, lipid‐binding and structural bioinformatics, we characterized five NtPLDδ isoforms. Distinct PLDδ isoforms preferentially localize to the cytoplasm or subapical PM. Using fluorescence recovery after photobleaching, domain deletion and swapping analyses we show that membrane‐bound PLDδs are tightly bound to PM, primarily via the central catalytic domain. Overexpression analyses suggested isoform PLDδ3 as the most important member of the PLDδ subfamily active in pollen tubes. Moreover, only PLDδ3 shows significant constitutive PLD activity in vivo and, in turn, PA promotes binding of PLDδ3 to the PM. This forms a positive feedback loop leading to PA accumulation and the formation of massive PM invaginations. Tightly controlled production of PA generated by PLDδ3 at the PM is important for maintaining the balance between various membrane trafficking processes that are crucial for plant cell tip growth. Significance Statement: Phosphatidic acid has been demonstrated as an important molecule defining the identity of plasma membrane, but its production from structural phospholipids by phospholipase D is still not completely understood. This study describes a comprehensive bioinformatic, localization and functional analysis of phospholipase Dδ subfamily in tobacco pollen tubes, showing that distinct isoforms have different membrane targeting mechanisms, dynamics, in vivo activity and the effect on cell membrane traffic. [ABSTRACT FROM AUTHOR]

Land plants with elaborated three-dimensional (3D) body plans have evolved from streptophyte algae. The streptophyte algae are known to exhibit varying degrees of morphological complexity, ranging from single-celled flagellates to branched macrophytic forms exhibiting tissue-like organization. In this review, I discuss mechanisms by which, during evolution, filamentous algae may have gained 2D and eventually 3D body plans. There are, in principle, two mechanisms by which an additional dimension may be added to an existing algal filament or cell layer: first, by tip growth-mediated branching. An example of this mechanism is the emergence and polar expansion of root hairs from land plants. The second possibility is the rotation of the cell division plane. In this case, the plane of the forthcoming cell division is rotated within the parental cell wall. This type of mechanism corresponds to the formative cell division seen in meristems of land plants. This literature review shows that of the extant streptophyte algae, the Charophyceae and Coleochaetophyceae are capable of performing both mechanisms, while the Zygnematophyceae (the actual sister to land plants) show tip growth-based branching only. I finally discuss how apical cells with two or three cutting faces, as found in mosses, may have evolved from algal ancestors. [ABSTRACT FROM AUTHOR]

Pollen tubes rapidly elongate, penetrate, and navigate through multiple female tissues to reach ovules for sperm delivery by utilizing a specialized form of polar growth known as tip growth. This process requires a battery of cellular activities differentially occurring at the apical growing region of the plasma membrane (PM), such as the differential cellular signaling involving calcium (Ca2+), phospholipids, and ROP-type Rho GTPases, fluctuation of ions and pH, exocytosis and endocytosis, and cell wall construction and remodeling. There is an emerging understanding of how at least some of these activities are coordinated and/or interconnected. The apical active ROP modulates exocytosis to the cell apex for PM and cell wall expansion differentially occurring at the tip. The differentiation of the cell wall involves at least the preferential distribution of deformable pectin polymers to the apex and non-deformable pectin polymers to the shank of pollen tubes, facilitating the apical cell expansion driven by high internal turgor pressure. Recent studies have generated inroads into how the ROP GTPase-based intracellular signaling is coordinated spatiotemporally with the external wall mechanics to maintain the tubular cell shape and how the apical cell wall mechanics are regulated to allow rapid tip growth while maintaining the cell wall integrity under the turgor pressure. Evidence suggests that exocytosis and endocytosis play crucial but distinct roles in this spatiotemporal coordination. In this review, we summarize recent advances in the regulation and coordination of the differential pectin distribution and the apical domain of active ROP by exocytosis and endocytosis in pollen tubes. [ABSTRACT FROM AUTHOR]