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Cell plate

The cell plate is a transient membranous structure that forms during cytokinesis in higher plant cells, serving as the precursor to the new cell wall that partitions the cytoplasm between two daughter cells.^[1] It arises from the fusion of Golgi-derived vesicles at the equatorial plane of the phragmoplast, a microtubule-based array that guides its centrifugal expansion toward the parental cell walls.^[2] This process ensures the physical separation of daughter cells while incorporating essential cell wall components, distinguishing plant division from the contractile ring mechanism in animal cells.^[3] The formation of the cell plate begins in late anaphase or telophase, shortly after chromosome segregation, with vesicles containing polysaccharides such as callose, pectins, hemicelluloses, and glycoproteins accumulating at the division site.^[1] These vesicles, derived from the trans-Golgi network, fuse to create a tubulo-vesicular network that matures through stages including a tubular network and a fenestrated sheet, ultimately fusing with the plasma membrane to complete cytokinesis.^[4] Key molecular players include SNARE proteins like KNOLLE for vesicle fusion, dynamin-related proteins for membrane remodeling, and tethering complexes such as the exocyst and TRAPPII for targeted delivery.^[2] Cellulose synthesis enzymes (CESAs) are recruited later to deposit microfibrils, replacing transient callose and strengthening the maturing wall.^[4] The cell plate's development is crucial for plant growth and tissue organization, as it establishes the plane of division and allows for the formation of plasmodesmata—channels enabling intercellular communication.^[1] Disruptions in this process, such as mutations in callose synthase genes like GSL8, can lead to incomplete walls and defective cell separation, highlighting its role in maintaining plant morphology.^[4] Studies emphasize the dynamic interplay of cytoskeletal elements, including actin filaments and microtubules, in directing vesicle trafficking and ensuring precise plate expansion.^[2] More recent research (2023–2025) has highlighted roles for lipids like phosphatidylinositol 4-phosphate (PI4P) in cell plate morphology transitions, endoplasmic reticulum (ER)-dependent membrane fusion in plasmodesmata formation, and proteins such as HYCCIN2 in recruiting the SH3P2-DRP1A complex for membrane tubulation.^[5]^[6]^[7]

Biological Context

Cytokinesis in Plants

Cytokinesis is the process by which the cytoplasm of a plant cell divides following mitosis, resulting in two distinct daughter cells. This physical separation ensures that each daughter cell receives a complete set of organelles and cytoplasmic components. In plants, cytokinesis is tightly coordinated with the later stages of mitosis, typically initiating during late anaphase and completing by the end of telophase, as the chromosomes decondense and nuclear envelopes reform around the daughter nuclei.^[1]^[8] Unlike animal cells, which employ a contractile ring of actin and myosin to constrict and furrow the plasma membrane, plant cells face unique challenges due to their rigid, preexisting cell walls composed primarily of cellulose and other polysaccharides. This inflexible extracellular matrix prevents the cell from pinching inward, necessitating an internal partitioning mechanism that builds a new cell wall from the inside out. The cell plate serves this critical role, forming as a transient structure that expands centrifugally to divide the cytoplasm and eventually fuses with the parental cell walls, thereby establishing a permanent septum between the daughter cells.^[1]^[4] The plane of division in plant cytokinesis is predetermined during the G2 phase of the cell cycle by the preprophase band, a cortical ring of microtubules and actin filaments that marks the future site where the cell plate will insert into the parental walls. This band ensures precise and oriented cell division, which is essential for proper tissue organization in plants. The phragmoplast, a microtubule array that forms during anaphase, guides the subsequent assembly and positioning of the cell plate along this predetermined plane.^[1]^[9]

Role of the Phragmoplast

The phragmoplast is a specialized cytoskeletal structure that emerges as a bipolar array of microtubules and actin filaments during the transition from anaphase to telophase in plant cytokinesis.^[10] This array forms between the separating daughter nuclei, originating from remnants of the central spindle, with microtubules oriented antiparallel and overlapping at their plus ends in the central midzone, while actin filaments contribute to the overall framework stability.^[10] The bipolar organization ensures a symmetrical scaffold that supports the initial assembly of the cell plate at the cell's equatorial plane.^[11] A primary function of the phragmoplast is to guide the delivery of Golgi-derived vesicles to the site of cell plate formation along microtubule tracks.^[10] Microtubule-based motor proteins, such as kinesins from the kinesin-12 family, transport these vesicles toward the midzone, where they accumulate and fuse to build the nascent cell plate.^[12] This directed trafficking is essential for concentrating vesicular materials precisely at the division site, preventing mislocalization during cytoplasmic partitioning.^[13] The positioning of the phragmoplast is predetermined by the preprophase band, a cortical array of microtubules and actin filaments that marks the future division plane during prophase.^[13] This band influences spindle orientation and directs the phragmoplast to expand toward the specified cortical site, ensuring accurate cell plate insertion and maintaining tissue organization.^[14] Phragmoplast dynamics involve centrifugal expansion, where the structure grows outward from the cell center to the periphery as the cell plate matures.^[10] This process is driven by microtubule polymerization at the leading edges and depolymerization at the trailing zones, coupled with kinesin-mediated sliding of antiparallel microtubules, resulting in an inside-out progression that mirrors the cell plate's extension.^[15]

Formation Process

Initiation and Vesicle Delivery

Cell plate formation initiates during telophase of mitosis, when the Golgi apparatus and trans-Golgi network (TGN) produce specialized vesicles laden with cell wall precursors, such as pectins and hemicelluloses, along with membrane components essential for partitioning the daughter cells. These cytokinetic vesicles, often termed cell plate vesicles, accumulate rapidly as the mitotic spindle transitions into the phragmoplast structure. The production is tightly regulated to ensure sufficient material delivery to the division site, marking the onset of de novo membrane assembly in plant cytokinesis.^[4] These vesicles are transported bidirectionally along phragmoplast microtubules, which serve as tracks guiding them toward the equatorial plane of the dividing cell. Microtubule plus ends at the equator facilitate vesicle capture and alignment, preventing random distribution and ensuring precise positioning for fusion. This directed transport, powered by motor proteins like kinesins, concentrates vesicles at the future cell plate site within minutes of telophase entry.^[4] Upon arrival, TGN-derived vesicles undergo initial docking and fusion events, coalescing to form a transient tubulo-vesicular network (TVN) that represents the earliest visible cell plate structure. This fusion stage, known as the fusion of vesicles stage (FVS), transitions into a tubulo-vesicular network (TVN), which further organizes into a more structured tubular network (TN) through homotypic membrane mergers. SNARE proteins play a central role in mediating these events; the cytokinesis-specific syntaxin KNOLLE (a Qa-SNARE) localizes to vesicle membranes and interacts with Qb/Qc-SNARE SNAP33 and R-SNAREs VAMP721/VAMP722 to drive docking and fusion specificity.^[16] These SNARE complexes ensure efficient, targeted membrane integration, preventing defects in cell plate assembly observed in mutants lacking these components.

Assembly and Expansion

Following initial formation, the cell plate expands centrifugally from the cell center toward the parental walls, a process orchestrated by the dynamic expansion of the phragmoplast, where microtubules reorganize from a dense disc into an expanding ring to direct vesicle delivery to the periphery.^[4] This outward growth ensures the cell plate contacts the existing cell walls, partitioning the cytoplasm into two daughter cells. The expansion involves the continuous addition and fusion of Golgi-derived vesicles at the leading edge of the cell plate, transforming it from a tubular-vesicular network into a fenestrated, sheet-like structure characterized by a honeycomb-like array of interconnected membranes.^[4] Dynamin-related proteins assist in membrane tubulation and remodeling during these fusion events, mediated by SNARE proteins such as syntaxins, which maintain the integrity and uniformity of the expanding membrane while allowing for the incorporation of additional material to fill gaps.^[4] Vesicle motility during this expansion phase relies on the actin cytoskeleton and myosin motors, particularly class VIII and XI myosins, which facilitate directed transport along actin filaments toward the phragmoplast equator and support the mechanical forces needed for membrane spreading.^[4]^[17] These actomyosin interactions ensure efficient vesicle delivery and prevent misalignment as the cell plate grows.^[17] The coordinated expansion is tightly regulated by signaling pathways, including the NPK1 MAP kinase kinase kinase, which is activated at the phragmoplast equator by kinesin-like proteins NACK1 and NACK2 to promote microtubule depolymerization.^[18] This kinase complex localizes precisely to drive lateral phragmoplast broadening, thereby enabling uniform cell plate growth without fragmentation.^[18]

Composition and Components

Vesicular Contents

The vesicles contributing to cell plate formation primarily originate from the trans-Golgi network (TGN), which serves as the main secretory compartment for delivering membrane and matrix materials during plant cytokinesis.^[19] Multivesicular bodies (MVBs) also contribute vesicles, providing additional endocytic recycling of components to support cell plate assembly.^[19] These vesicles are guided to the division site by the phragmoplast, ensuring targeted fusion and deposition.^[20] Membrane components within these vesicles include phospholipids that form the lipid bilayers of the nascent cell plate, which mature into the new plasma membranes separating daughter cells.^[21] Integral membrane proteins, such as those involved in vesicle fusion and membrane stabilization, are also transported to establish the functional boundaries of the dividing cells.^[19] Cell wall precursors constitute a major cargo of TGN-derived vesicles, including pectins that provide gel-like matrix for initial structural integrity; for instance, demethylesterified pectins are delivered alongside pectin methylesterases (e.g., AtPME1) that modulate pectin cross-linking post-deposition.^[19] Hemicelluloses, such as xyloglucans, are synthesized in the Golgi and trafficked via TGN vesicles to integrate with the forming cell wall matrix.^[19] Glycoproteins, including extensins and arabinogalactan proteins, are also delivered via these vesicles to contribute to cell wall structure and intercellular communication.^[4] Cellulose synthases (CESAs), assembled in the Golgi apparatus, are delivered through TGN/SYP61-defined vesicles to the cell plate, where they initiate cellulose microfibril synthesis for long-term wall reinforcement.^[22] A temporary matrix is established by callose (β-1,3-glucan), which provides structural support during cell plate expansion and stabilization before its replacement by permanent wall components; callose synthases, such as AtGSL8, localize to the forming cell plate to synthesize this polymer in situ, potentially incorporating precursors or complex subunits delivered via TGN vesicles.^[23]

Key Proteins and Regulators

Phragmoplastin, a dynamin-like GTPase, plays a central role in cell plate construction by facilitating the fission and fusion of vesicles at the developing plate during cytokinesis in plants. This protein, identified in soybean and conserved across plant species, localizes to the cell plate and associates with exocytic vesicles that deliver materials such as pectins, promoting membrane remodeling essential for plate assembly. Overexpression studies have shown that excess phragmoplastin leads to callose accumulation and growth arrest, underscoring its precise regulation for proper vesicle dynamics.^[24]^[25] The syntaxin family, particularly the plant-specific KNOLLE (SYP111), is crucial for vesicle docking and fusion at the cell plate. KNOLLE, a cytokinesis-specific syntaxin, accumulates at the division plane during late mitosis and mediates the targeted fusion of Golgi-derived vesicles, ensuring accurate partitioning of cytoplasm. Other members of the syntaxin of plants (SYP) family, such as those in the SYP1 group, contribute to post-Golgi trafficking that supports cell plate formation, though KNOLLE's role is uniquely restricted to dividing cells. Mutations in KNOLLE result in defective cell plates and multinucleate cells, highlighting its indispensability.^[26] Rab GTPases regulate vesicle trafficking to the cell plate by coordinating the recruitment and delivery of secretory vesicles from the trans-Golgi network. Members like RABA1e and RABA5c localize to the cell plate edges, directing polarized transport and fusion events during plate expansion. These small GTPases interact with tethering factors to position vesicles accurately, preventing mislocalization that could disrupt cytokinesis.^[27]^[28] Tethering complexes, including TRAPPII and the exocyst, play essential roles in vesicle targeting and fusion during cell plate assembly and maturation. The TRAPPII complex is required for early biogenesis of the cell plate, facilitating vesicle fusion throughout cytokinesis, while the exocyst complex acts later to promote expansion and integration with parental walls. These complexes work sequentially with Rab GTPases and SNAREs to ensure precise membrane delivery.^[4]^[29] Dynamin-related proteins (DRPs), beyond phragmoplastin, contribute to membrane remodeling at the cell plate through GTPase activity that drives tubulation and scission. Plant DRP1 and DRP2 families localize to the division site, aiding in the reshaping of fused membranes into a planar structure. Cytoskeletal motors, including kinesin-14 proteins such as ATK5, organize microtubules to guide vesicle delivery, while myosin VIII associates with microtubule ends to facilitate actin-myosin-based transport of vesicles toward the plate. These motors ensure directed movement, with kinesin-14 promoting phragmoplast expansion and myosin VIII supporting rapid trafficking under high turgor pressure.^[30]^[31]^[32]

Maturation and Integration

Cell Wall Development

Following the initial formation of a matrix primarily composed of callose and pectins delivered via vesicles, the cell plate undergoes maturation into a permanent cell wall through the deposition of structural polysaccharides. Cellulose microfibrils are synthesized and deposited by rosette-shaped cellulose synthase complexes (CSCs), which consist of multiple cellulose synthase catalytic subunits (CESAs) organized into hexameric rosettes in the maturing cell plate membrane.^[33] In Arabidopsis, the cellulose synthase-like D5 (CSLD5) protein plays a critical role in this process, providing essential β-1,4-glucan synthase activity specifically during cell plate formation to reinforce the structure.^[34] This deposition begins at the late tubular network (TN) stage and continues through the planar fenestrated sheet (PFS) stage, aligning with the cell plate's flattening and increased rigidity.^[4] Remodeling of the transient callose matrix is essential for transitioning to a durable wall, involving the degradation of callose and subsequent cross-linking of matrix components. Callose, a β-1,3-glucan, accumulates transiently during early cell plate assembly but is actively removed by hydrolytic enzymes such as β-1,3-glucanases once the plate stabilizes, preventing excessive rigidity and allowing for further maturation.^[4] Recent studies (as of 2025) indicate that phosphatidylinositides, such as PI4P and PI(4,5)P2, regulate the morphology transition during this remodeling by controlling lipid asymmetry and membrane curvature, while also modulating callose biosynthesis to prevent excessive deposition.^[35] Concurrently, pectins—initially deposited as methyl-esterified forms from Golgi-derived vesicles—are de-esterified by pectin methylesterases, enabling calcium-mediated cross-linking that forms a cohesive gel-like matrix and enhances wall integrity.^[4] The synthesis of middle lamella components occurs simultaneously to ensure adhesion between daughter cells, with homogalacturonan pectins cross-linked by calcium ions to form calcium pectates, the primary constituents of this intercellular layer. Rhamnogalacturonan II (RG-II), another complex pectin, is also incorporated at the cell plate to facilitate borate-mediated dimerization and further stabilize the structure.^[4] These processes unfold during the late expansion phase of cell plate development, particularly in the PFS stage, guaranteeing structural support and proper cell separation before fusion with parental walls.^[4]

Fusion with Parental Walls

During the final phase of plant cytokinesis, the cell plate undergoes centrifugal maturation, wherein its margins progressively anneal with the parental plasma membranes and cell walls, establishing a continuous boundary between daughter cells. This process begins as the expanding cell plate, guided by the phragmoplast, contacts the equatorial periphery of the mother cell, ensuring precise alignment and integration without disrupting existing cellular structures. Recent research (as of 2025) highlights the essential role of endoplasmic reticulum (ER)-plasma membrane (PM) contact sites, regulated by VAP27 proteins, in recruiting the actin cytoskeleton via the SCAR/WAVE complex to facilitate cell plate expansion and proper orientation during integration.^[36]^[37]^[38] Membrane fusion at these margins is orchestrated by the exocyst complex, which tethers incoming vesicles to the target sites on the parental plasma membrane, thereby promoting spatial accuracy prior to SNARE-mediated merger. In Arabidopsis thaliana, exocyst subunits such as EXO84b localize to the cell plate margins and facilitate vesicle docking, while SEC6 directly interacts with the SNARE regulator KEULE to coordinate fusion events. Complementary SNARE proteins, including R-SNAREs VAMP721 and VAMP722, drive the actual membrane fusion; mutants lacking these proteins display retarded cell plate expansion, binucleate cells, and incomplete attachment to parental walls, underscoring their essential role.^[39]^[40]^[16] To achieve wall continuity, a shared polysaccharide matrix is deposited at the fusion junctions, sealing the nascent cell plate to the parental walls and transforming the transient structure into a rigid, permanent barrier. This involves the targeted delivery of callose for initial stabilization and cellulose along with hemicelluloses like xyloglucan for long-term reinforcement, ensuring seamless integration without gaps or fenestrae.^[4]^[41] Full fusion triggers completion signals that initiate phragmoplast disassembly, with microtubules depolymerizing outward from the center as the division plane stabilizes. This regulated breakdown, mediated by severing proteins like KATANIN1 and phosphoinositide signaling, halts further vesicle trafficking and marks the end of cytokinesis.^[15]^[42]

Comparisons and Variations

With Animal Cytokinesis

In animal cells, cytokinesis occurs through the formation of a contractile ring composed of actin filaments and myosin II motors, which assembles at the equatorial plane and constricts the plasma membrane inward to create a cleavage furrow that pinches the cell into two daughter cells.^[1] This mechanism relies on the deformability of the flexible plasma membrane and cytoskeleton to physically bisect the cell. In contrast, plant cytokinesis via cell plate formation builds a new partition internally from Golgi-derived vesicles, necessitated by the rigid cell wall that prevents furrow ingression. While animal cells deform the existing plasma membrane through contraction, plant cells expand the cell plate centrifugally to fuse with the parental walls, generating a new plasma membrane and cell wall without relying on a contractile apparatus.^[43] The phragmoplast, a microtubule array guiding vesicle delivery, is absent in animals, highlighting the structural adaptations for walled cells. Despite these differences, both processes share conserved regulatory elements, including Rho GTPases, which determine the division plane and coordinate cytokinesis with mitosis. In animals, RhoA activation at the equator promotes contractile ring assembly and furrow progression.^[44] In plants, ROP (Rho of plants) GTPases similarly regulate vesicle trafficking and cytoskeletal dynamics to position and expand the cell plate.^[45] The cell plate mechanism evolved as an adaptation for cytokinesis in walled cells, originating in streptophyte green algae and conserved in land plants to accommodate turgor pressure and cell wall rigidity.^[46] This contrasts with the ancestral cleavage furrow-like division in unwalled eukaryotes, underscoring the divergence driven by cell wall acquisition in the plant lineage.^[47]

Across Plant Lineages

Cell plate formation in angiosperms follows the canonical phragmoplast-guided process, where Golgi-derived vesicles accumulate at the division site and fuse centrifugally to form a planar structure that matures into the new cell wall, incorporating complex polysaccharides such as cellulose, hemicelluloses, and pectins for structural integrity.^[48] This mechanism ensures precise partitioning during somatic and meiotic divisions, with the phragmoplast providing microtubule-based guidance for vesicle delivery. In gymnosperms, such as conifers, cell plate assembly is similarly phragmoplast-dependent and centrifugal, relying on Golgi-derived vesicles for initial fusion, but exhibits subtle variations in wall composition, including higher incorporation of glucomannan and lignin precursors that contribute to more rigid secondary wall development post-fusion.^[48]^[49] Ferns, as pteridophytes, also employ a phragmoplast-guided centrifugal process with Golgi vesicles, though their cell plates often integrate distinct polysaccharides like higher levels of arabinoxylans and lignin-related compounds compared to angiosperms, reflecting adaptations to terrestrial vascularity.^[48]^[50] Advanced streptophyte green algae, such as Coleochaete and Chara, exhibit phragmoplast-guided cell plate formation with centrifugal expansion, though with simpler structures compared to land plants, resulting in cellulose-based septa without the multilayered complexity of land plant walls.^[48]^[47] Across streptophytes, the core vesicle fusion machinery, including conserved SNARE proteins and phragmoplastin, remains evolutionarily stable from algal ancestors to land plants, enabling the transition from aquatic septation to robust terrestrial cytokinesis despite lineage-specific adaptations.^[51]^[47]

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