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Synaptonemal complex

The synaptonemal complex (SC) is a highly conserved, meiosis-specific multiprotein structure that assembles between paired homologous chromosomes during the prophase of the first meiotic division, facilitating their close alignment and synapsis.^[1] This zipper-like assembly, first observed in the 1950s, spans approximately 100 nm between chromosomes and is essential for the proper execution of meiotic recombination.^[2] Composed of a tripartite architecture, the SC consists of two lateral elements along each chromosome axis, transverse filaments connecting the homologs, and a central element in between.^[3] Key protein components of the SC include SYCP2 and SYCP3, which form the lateral elements and provide structural support akin to axial elements during pre-synapsis; SYCP1, which constitutes the transverse filaments and mediates chromosome pairing; and central element proteins such as SYCE1, SYCE2, SYCE3, TEX12, and SIX6OS1.^[1] Additional elements, including meiosis-specific cohesins (e.g., REC8, STAG3) and HORMAD proteins (HORMAD1, HORMAD2), regulate assembly and chromatin interactions.^[1] The SC's formation is tightly coordinated with chromatin condensation and the initiation of double-strand breaks by SPO11, ensuring homologous recombination proceeds accurately.^[2] Functionally, the SC stabilizes homologous chromosome pairing, promotes the maturation of double-strand breaks into crossovers, and enforces crossover interference to distribute recombination events evenly along chromosomes, thereby preventing nondisjunction and aneuploidy during gamete formation.^[3] In many organisms, including mammals, the SC exhibits liquid-crystalline properties that drive its polymerization and alignment, with self-assembly mechanisms involving multivalent protein interactions.^[3] Disruptions in SC components, such as mutations in SYCP3 or SYCE1, lead to synaptic failure, reduced fertility, and associations with conditions like non-obstructive azoospermia or premature ovarian insufficiency in humans.^[1] Across sexually reproducing species—from yeast (e.g., Zip1) to flies (e.g., C(3)G) and mammals—the SC's core role in meiosis remains invariant, underscoring its evolutionary significance.^[3]

Overview and Discovery

Definition and Basic Structure

The synaptonemal complex (SC) is a meiosis-specific, proteinaceous structure that forms between homologous chromosomes during prophase I of meiosis, serving as a scaffold to mediate their pairing and alignment.^[4] It plays a crucial role in facilitating genetic recombination and ensuring the accurate segregation of chromosomes by stabilizing synapsis along the full length of the paired homologs. The SC exhibits a highly conserved tripartite architecture, consisting of two parallel lateral elements (LEs) that run along the axes of sister chromatids, a central element (CE) positioned in the middle, and transverse filaments (TFs) that bridge the LEs to the CE.^[4] This ladder-like configuration aligns the homologous chromosomes in close proximity, with the LEs providing attachment points for chromatin loops and the TFs and CE forming the interconnecting core. Under electron microscopy, the SC appears as a distinctive ladder-like structure, with transverse filaments visible as closely spaced crossbars forming a periodic appearance along the length of the complex. The overall width of the SC, measured as the distance between the two LEs, is approximately 100-200 nm, while the central region spans about 100 nm, and the structure extends continuously along the entire length of the paired chromosomes, which can reach several micrometers in total. Formation of the SC is a prerequisite for proper chromosome segregation in meiosis, as its absence leads to meiotic defects and aneuploidy.^[4]

Historical Context and Key Discoveries

The synaptonemal complex (SC) was first observed in the mid-1950s through advancements in electron microscopy, which allowed visualization of fine chromosomal structures during meiotic prophase. In 1956, Montrose J. Moses reported the presence of a novel ribbon-like chromosomal component in primary spermatocytes of the crayfish Procambarus clarkii, describing it as a continuous structure approximately 100 nm wide running parallel to the chromosomes along their length. Independently in the same year, Don W. Fawcett identified similar structures in spermatocytes of vertebrates, including the pigeon (Columba livia) and cat (Felis catus), noting their tripartite organization consisting of two dense lateral elements flanking a less dense central region. These early observations established the SC as a conserved feature of meiosis across invertebrates and vertebrates, though its functional significance remained unclear at the time. The term "synaptonemal complex" was introduced in the early 1960s to describe this structure, drawing from its role in synaptic (pairing) processes and its nemaline (thread-like) appearance under electron microscopy. Pioneering studies by Theodore F. Roth in 1966 further characterized SC dynamics in mosquito oocytes, revealing changes during meiotic prophase and suggesting involvement in chromosome pairing. Concurrently, David von Wettstein and colleagues advanced the field through detailed ultrastructural analyses in fungi and plants, such as Neottiella and Lilium, demonstrating the SC's zipper-like assembly between homologous chromosomes. By the late 1960s, Moses synthesized these findings in a comprehensive review, emphasizing the SC's universal occurrence in sexually reproducing eukaryotes and its association with pachytene-stage bivalents.^[5] Key breakthroughs in the 1970s and 1980s linked the SC to meiotic recombination and segregation. Von Wettstein's group, using serial section reconstructions, proposed that SC formation facilitates genetic exchange by stabilizing homologous pairing, a model supported by observations of recombination nodules along the central region in Bombyx mori. In 1984, von Wettstein, Stig Rasmussen, and Peter Holm published a seminal review integrating cytological and genetic data, arguing that the SC provides the structural framework for crossover interference and proper chromosome disjunction. These studies shifted the perception of the SC from a mere morphological artifact to an essential regulator of meiotic fidelity, paving the way for molecular identifications in the 1990s. The 50th anniversary of its discovery was marked in 2006, highlighting ongoing research into its conserved role across species.^[6]^[7]

Molecular Composition

Core Structural Proteins

The synaptonemal complex (SC) features a tripartite architecture comprising lateral elements, transverse filaments, and a central element, with core structural proteins providing the essential scaffold. The lateral elements (LEs) are primarily composed of synaptonemal complex protein 2 (SYCP2) and synaptonemal complex protein 3 (SYCP3), which form elongated, fibrous structures along the lengths of homologous chromosomes.^[8] SYCP3 consists of extended coiled-coil domains that enable it to polymerize into tetrameric filaments approximately 20 nm long, serving as a structural backbone for chromosome axis organization.^[9] SYCP2, also featuring coiled-coil motifs, integrates with SYCP3 to enhance the rigidity and stability of the LEs through heterodimer formation, where the C-terminal region of SYCP2 directly binds SYCP3, facilitating their co-polymerization along chromatin.^[10] These SYCP2-SYCP3 heterodimers provide mechanical support, maintaining the linear arrangement of looped chromatin domains during meiosis.^[11] The transverse filaments (TFs) are predominantly formed by synaptonemal complex protein 1 (SYCP1), a large coiled-coil protein that acts as the primary architect of the SC's zipper-like lattice, while the central element (CE) is mainly composed of proteins such as SYCE1, SYCE2, SYCE3, and TEX12 that assemble with the C-terminal domains of SYCP1. SYCP1 molecules self-assemble in a head-to-head and tail-to-tail manner, with their central coiled-coil domains spanning the ~100 nm gap between paired LEs to form the TFs.^[12] The N-terminal domains of SYCP1 are globular and mediate attachment to the LEs by binding directly to DNA and interacting with SYCP2/SYCP3 components, thereby anchoring the homologous chromosomes in close proximity.^[13] At the opposite end, the C-terminal knobs of SYCP1 protrude into the CE, where they oligomerize with SYCE proteins to create a periodic lattice that stabilizes the overall SC structure and enforces precise alignment.^[12] This self-assembly mechanism of SYCP1 ensures the supramolecular organization essential for synapsis.^[13] Key interactions among these proteins reinforce the SC scaffold: SYCP3-SYCP2 heterodimers confer structural rigidity to the LEs, while the C-terminal domain of SYCP2 also bridges to SYCP1, linking the LEs to the TFs and CE.^[11] SYCP1's self-assembly into the CE is driven by hydrophobic interactions and coiled-coil associations, independent of other components in vitro.^[12] Post-translational modifications, such as phosphorylation of SYCP3 at specific serine residues, regulate the stability of these protein complexes by modulating polymerization dynamics and preventing premature disassembly.^[14] Atomic models derived from structural studies have elucidated these architectures. X-ray crystallography has provided insights into SYCP3's tetrameric organization at near-atomic resolution (2.2 Å), revealing its extended helical bundles.^[9] For SYCP1, X-ray crystallography of its coiled-coil domains achieved resolutions around 2.9 Å, detailing the molecular basis of TF and CE formation, while cryo-EM reconstructions at approximately 10 Å confirmed the in situ lattice arrangement.^[12] These models highlight the precise intermolecular contacts that underpin SC integrity.^[15]

Accessory and Regulatory Proteins

The synaptonemal complex (SC) relies on accessory and regulatory proteins to modulate its assembly, stability, and integration with meiotic recombination processes, without constituting the primary structural scaffold. These proteins facilitate interactions between the SC and chromosome axes, enforce crossover specificity, and monitor synapsis fidelity, ensuring proper homologous chromosome pairing and genetic exchange. Meiotic cohesins, including REC8-containing complexes, interact with the chromosome axes organized by the core proteins to maintain sister chromatid cohesion and position recombination sites. These associations are essential for linear axis compaction and facilitating SC attachment during leptonema.^[16] Crossover regulators, notably the MLH1-MLH3 (MutLγ) endonuclease complex, associate with the SC to direct the resolution of double Holliday junctions into crossovers. This complex localizes to SC sites marked for interference-sensitive crossovers, where it incises joint molecules in a PCNA- and EXO1-dependent manner, ensuring biased resolution within the SC environment.^[17]^[18] Disruption of MLH1-MLH3 leads to reduced crossover numbers and impaired double-strand break repair, highlighting its role in channeling recombination outcomes.^[19] Kinases like ATM and ATR phosphorylate SC-associated proteins to coordinate recombination progression and DNA damage responses. ATM/ATR target sites on central region components, such as SYP-4 in C. elegans (a homolog of mammalian SYCE proteins), to promote homologous recombination over non-homologous end joining and signal repair completion.01403-8) In mice, ATR activation on unsynapsed axes regulates DSB processing, while ATM modulates cohesin dynamics to support axis integrity during prophase I.^[20]^[21] These phosphorylation events provide feedback to prevent progression until recombination is resolved. HORMAD1 and HORMAD2 function as anti-synapsis sentinels, localizing to unsynapsed chromosome axes to inhibit non-homologous pairing and promote DSB formation via recruitment of SPO11 accessories. HORMAD1/2 are depleted from synapsed regions by TRIP13 ATPase, allowing SC polymerization only between homologs; their persistence on axes activates checkpoints that eliminate defective meiocytes.^[22]^[23] In mice, HORMAD2 further recruits ATR to amplify signaling against asynapsis.^[24] Mutations in these accessory proteins often result in SC instability and meiotic failure. For instance, Hormad1 knockout in mice disrupts axis loading of HORMAD2, leading to widespread asynapsis, reduced DSBs, and complete infertility due to checkpoint-mediated oocyte elimination.^[22] Similarly, deficiencies in MLH3 or ATR compromise crossover formation and axis maintenance, causing aneuploidy and sterility.^[20]^[17] These effects emphasize the regulatory proteins' necessity for SC integrity and meiotic fidelity.

Assembly and Dynamics

Stages of Formation

The formation of the synaptonemal complex (SC) occurs progressively during meiotic prophase I, beginning with the establishment of chromosome axes and culminating in the full stabilization of homologous chromosome pairing. In the pre-leptotene stage, following premeiotic DNA replication, cohesin complexes such as those containing REC8, SMC1β, and STAG3 load along the chromosomes to form the initial linear scaffold known as axial elements (AEs).^[25] These AEs are further reinforced by the loading of structural proteins SYCP2 and SYCP3, which polymerize to create a rigid, proteinaceous core that organizes chromatin into loops and prepares the chromosomes for alignment; this process is essential for subsequent synapsis and requires intact cohesin function.^[25]^[26] Transitioning into leptotene and zygotene, double-strand breaks (DSBs) induced by SPO11 facilitate the search for homologous partners, while synapsis initiates primarily at chromosome telomeres in many organisms, forming a clustered "bouquet" configuration that promotes efficient pairing.^[27] In this phase, the lateral elements (LEs), composed of SYCP2 and SYCP3, elongate and align in parallel between homologs, with transverse filament protein SYCP1 beginning to polymerize between them to bridge the AEs and initiate SC zipper-like extension from the initiation sites.^[27]^[28] DSBs play a key role in guiding this parallel synapsis, stabilizing early pairing contacts along the chromosome arms. By the pachytene stage, synapsis achieves full closure along the length of each bivalent, completing the SC structure with the maturation of the central element (CE). Proteins such as SYCE1, SYCE2, SYCE3, and TEX12 assemble within the CE to cross-link and rigidify the SYCP1 transverse filaments, ensuring stable homolog association across the entire chromosome. This zipper-like progression transforms the partial alignments of zygotene into a continuous tripartite complex, marking the completion of SC assembly.^[25] In model organisms like mice, SC formation unfolds over a defined timeline during spermatogenesis: pre-leptotene axis loading occurs around postnatal days (PND) 7-8, leptotene/zygotene synapsis initiation around PND 10-12, and pachytene progression from PND 12-18 in the first synchronous wave of germ cells.^[29] Progression through these stages is monitored by checkpoints, including the synapsis checkpoint mediated by the ATM kinase, which detects incomplete pairing or unrepaired DSBs along unsynapsed axes and activates apoptosis to eliminate defective meiocytes, thereby safeguarding genome integrity.^[30] Interspecies variations highlight the flexibility of SC assembly mechanisms; for instance, mammals typically initiate synapsis telomerically via the bouquet arrangement to facilitate rapid pairing in large genomes, whereas in Caenorhabditis elegans, initiation occurs interstitially at multiple sites without telomere clustering, relying instead on chromosome movement and DSB distribution for progressive SC polymerization.^[25]^[31] These differences reflect adaptations to genome size and nuclear organization but converge on the core principle of sequential, regulated assembly to ensure accurate homolog recognition.^[31]

Mechanisms of Disassembly

The disassembly of the synaptonemal complex (SC) initiates at the onset of diplotene during meiotic prophase I, marking the transition from pachytene and enabling homologous chromosomes to separate while preserving chiasmata for proper segregation. This process is tightly regulated by post-translational modifications, including phosphorylation and dephosphorylation events that loosen SC structural integrity. For instance, protein phosphatase 2A (PP2A), particularly the B56 subunit, plays a role in modulating SC dynamics; its overexpression in Drosophila leads to premature disassembly by counteracting stabilizing phosphorylations, highlighting its potential involvement in dephosphorylating key lateral element proteins like SYCP3 to initiate loosening.^[32] In mammals, disassembly often begins with a gradual release at telomeric regions, where the SC opens from chromosome ends, allowing desynapsis to proceed inward while remnants persist as short loops or fragments along non-sister chromatid arms. This telomeric opening facilitates the resolution of interlocks and entanglements, with SC components like SYCP1 and SYCP3 initially retained at centromeres before full dispersal. Live-cell microscopy in budding yeast (Saccharomyces cerevisiae) has visualized this dynamic dissolution, showing that Zip1, the central element protein analogous to mammalian SYCP1, disassembles monophasically at a rate of 66 ± 30 nm/min starting from chromosome ends, ensuring ordered progression without disrupting axis organization.^[33]^[34] Proteolytic degradation via the ubiquitin-proteasome system further drives SC breakdown, targeting core proteins for removal after recombination completion. The anaphase-promoting complex/cyclosome (APC/C), activated by its co-activator FZR1 (also known as CDH1), ubiquitinates substrates to promote timely disassembly; in mice, FZR1 deficiency allows initial SYCP3 loading but arrests progression beyond zygotene, indirectly implicating APC/C in clearing SC components like SYCP1 and SYCP3 to prevent persistence. In mammalian oocytes, rising cyclic AMP (cAMP) levels around birth signal this degradation, as inhibiting cAMP synthesis with MDL-12,330A disrupts SYCP1 disassembly, leading to prolonged linear staining indicative of stalled desynapsis.^[35]^[36] Failure in SC disassembly can result in persistent structures that trigger meiotic checkpoints and arrest. In S. cerevisiae spo11 mutants, which lack double-strand breaks essential for recombination, SC formation occurs at low frequency (∼10% in pachytene-arrested double mutants with ndt80Δ), but the inability to resolve these non-productive complexes leads to near-complete spore inviability (0.2%) due to unresolved pairing defects. Such persistence underscores the SC's role as a checkpoint sensor, where incomplete disassembly halts progression to metaphase I, ensuring only properly recombined chromosomes advance.^[37]

Functions in Meiosis

Chromosome Synapsis and Pairing

Chromosome synapsis refers to the intimate, stable pairing of homologous chromosomes along their entire length during meiotic prophase I, a process mediated by the synaptonemal complex (SC) scaffold that physically links the chromosome axes.^[3] This pairing ensures precise alignment, collinear organization, and subsequent genetic exchanges between homologs. The SC acts as a molecular zipper, polymerizing between the lateral elements of paired chromosomes to stabilize this association once initial contacts are established.^[3] Homolog recognition precedes full SC assembly and often involves the bouquet stage, where telomeres cluster at the nuclear envelope to facilitate initial search and alignment.^[38] In this configuration, dynamic chromosome movements, driven by microtubule attachments via the LINC complex, promote homologous interactions by reducing the search space and enhancing collision rates between matching sequences.^[38] Telomere clustering, observed across eukaryotes like fission yeast and mammals, initiates at the leptotene-zygotene transition and disperses as synapsis progresses, setting the stage for SC-mediated stabilization.^[38] The SC enforces collinearity by rigidly aligning homologous axes and preventing ectopic pairing with non-homologous chromosomes, a process regulated by HORMAD proteins such as Hop1 in yeast or HORMAD1/2 in mammals.^[39] These proteins bind unsynapsed chromatin via their HORMA domains and chromatin-binding regions, promoting homology-directed interactions while inhibiting off-target associations; for instance, mutations in Hop1's loop2 region lead to ectopic HORMAD accumulation and defective synapsis.^[39] Additionally, SC-associated chromatin loops, anchored by cohesin-based axes, organize higher-order domains that further support aligned pairing, with loop extrusion models indicating that these structures maintain ~100 nm inter-homolog distances for stable synapsis.^[40] Experimental evidence from fluorescence in situ hybridization (FISH) in SC mutants underscores the SC's role in sustaining homolog proximity. In Caenorhabditis elegans syp-2 mutants, which lack SC assembly, FISH probes reveal initial homolog pairing at chromosome ends but a sharp decline in alignment by late pachytene, contrasting with wild-type nuclei where SC maintains close juxtaposition throughout.^[41] Similar FISH analyses in other systems confirm that SC disruption results in dispersed homolog signals, highlighting its necessity for persistent physical closeness.^[41] As a homology sensor, the SC excludes non-homologous chromosomes from stable pairing, thereby preventing segregation errors like nondisjunction and aneuploidy.^[3] This quality control is evident in heterosynapsis assays, where SC forms between non-homologs only under forced conditions, but normally prioritizes homology to ensure accurate meiotic progression and reduce genetic instability.^[3]

Role in Genetic Recombination

The synaptonemal complex (SC) is essential for facilitating genetic recombination during meiosis by stabilizing the repair of programmed double-strand breaks (DSBs) induced by the Spo11 protein. Spo11 initiates DSB formation on chromatin during the leptotene stage, creating free DNA ends that invade homologous chromosomes for repair via homologous recombination; the SC, assembling along paired chromosomes, provides a structural scaffold that stabilizes these invaded strands and promotes strand invasion and exchange between homologs.^[42]^[43] The SC also acts as a platform for crossover interference, a process that ensures even spacing of crossovers along chromosomes, with MutLγ complex components like MLH1 forming discrete foci on the SC central region to designate sites of future class I crossovers. These MLH1 foci correlate directly with chiasmata positions and are influenced by SC integrity, as disruptions in SC proteins lead to altered focus distribution and reduced interference.^[44]^[45] In budding yeast, ZMM proteins, including Zip1 (a functional homolog of mammalian SYCP1) and Zip3, interact with the SC to bias DSB repair toward class I crossovers, promoting the formation of interference-sensitive recombination events while suppressing non-crossovers. Zip1 polymerizes to elongate the SC transverse filaments, stabilizing designated crossover sites, whereas Zip3 acts as a nucleation factor for SC assembly at early recombination intermediates.^[46]^[47] The SC enforces crossover assurance, ensuring that each homologous chromosome pair receives at least one crossover to secure proper segregation, a function mediated through its coordination of recombination progression and feedback mechanisms that prevent aneuploidy.^[48] Mechanistically, the beam-film model describes how the SC imposes physical stress along chromosome axes, akin to a beam under tension overlaid by a contracting film, which regulates strand exchange by propagating stress relief from initial crossovers to inhibit nearby events and promote distal ones.^[49] Genetic studies in SC mutants provide evidence for these roles; for instance, Sycp3-/- mice lacking a key lateral element protein exhibit reduced chiasma numbers and impaired crossover formation, with surviving oocytes showing an average of ~21 MLH1 foci per cell (compared to ~28 in wild-type) despite DSB repair, underscoring the SC's necessity for efficient recombination outcome.^[50]

Evolutionary and Cellular Necessity

Conservation Across Eukaryotes

The synaptonemal complex (SC) is a ubiquitous structure in meiotic cells across diverse eukaryotic lineages, including fungi, plants, animals, and protists, underscoring its fundamental role in sexual reproduction.^[51] This conservation extends to all major eukaryotic clades, where the SC facilitates the intimate pairing of homologous chromosomes during prophase I of meiosis.^[52] Its presence in such a broad phylogenetic distribution highlights the SC as a hallmark of meiosis, essential for stabilizing chromosome interactions and promoting genetic recombination.^[53] At the molecular level, core SC proteins exhibit remarkable conservation, with orthologs of the mammalian transverse filament protein SYCP1 and lateral element protein SYCP3 identified throughout eukaryotes. For instance, Zip1 in the fungus Saccharomyces cerevisiae functions as the primary transverse filament component, analogous to SYCP1, while SYP-1 in the nematode Caenorhabditis elegans serves as a direct ortholog of SYCP1, mediating central region assembly.^[54] Similarly, SYCP3 orthologs, such as Hop1 in yeast, form the lateral elements that anchor chromosomes to the SC scaffold. These orthologs share structural motifs, including coiled-coil domains, that enable the zipper-like polymerization essential for synapsis, demonstrating functional equivalence despite sequence divergence.^[55] The SC's functional universality is evident in its requirement for chromosome synapsis in nearly all studied eukaryotes, where its absence disrupts meiotic progression and leads to infertility or aneuploidy.^[45] Phylogenetic evidence points to an ancient origin of the SC, likely in the last eukaryotic common ancestor around 1.8 billion years ago, predating the diversification of eukaryotic supergroups and coinciding with the evolution of meiosis itself.^[56] Detailed investigations in model organisms have reinforced this conservation: in S. cerevisiae, Zip1-driven SC formation ensures crossover control; in the plant Arabidopsis thaliana, central element proteins like SCEP3 interlink synapsis initiation with recombination; and in the mouse Mus musculus, SYCP1 and SYCP3 coordinate full-length homolog pairing.90114-6)^[57]^[58] In contrast, the SC is absent in certain asexual lineages, including some parthenogenetic species that have secondarily lost meiosis, such as specific dinoflagellates where key SC-encoding genes are missing, reflecting the structure's tight linkage to sexual reproduction.^[59] This loss underscores the SC's evolutionary indispensability for meiotic fidelity in sexually reproducing eukaryotes.^[60]

Exceptions and Variations

In male Drosophila melanogaster, the synaptonemal complex (SC) is entirely absent during meiosis, which proceeds without recombination in an achiasmate manner.^[61] Homologous chromosomes pair instead through heterochromatic regions and specialized proteins like MNM and SNM, ensuring segregation without SC-mediated synapsis.^[62] This exception highlights an evolutionary adaptation for male-specific meiotic fidelity in the absence of crossover formation.^[63] In Caenorhabditis elegans, the SC exhibits a modified architecture compared to the ladder-like structure in mammals, forming as continuous ribbon-like sheets composed of SYP proteins that polymerize uniformly between paired chromosomes.^[64] These sheets lack discrete transverse filaments, instead providing broad adhesion along the entire length of homologs to facilitate recombination.^[65] This sheet-like organization supports efficient crossover control in the nematode's compact genome.^[52] In polyploid plants, such as allotetraploid Brassica napus or hexaploid Avena sativa, multiple SCs form per nucleus to accommodate the increased number of homologous or homeologous chromosomes. SC initiation varies, often occurring at multiple distal or interstitial sites per bivalent, contrasting with the more uniform zipper-like progression in diploids, which aids in stabilizing multivalent pairings during meiosis.^[66] These adaptations mitigate aneuploidy risks in polyploids by promoting balanced segregation.^[67] Among fungi, Ustilago maydis represents an exception where no SC forms despite functional meiotic recombination.^[68] Electron microscopy and genomic analyses confirm the absence of canonical SC components, yet homologous chromosomes pair and recombine effectively, suggesting alternative mechanisms like direct protein interactions suffice for genome reduction in this basidiomycete.^[69] This partial reliance on recombination without full SC assembly underscores flexibility in fungal meiosis.^[70] Evolutionary loss or modification of the SC correlates with sex chromosome differentiation, as seen in birds where the ZW pair forms only partial SCs in the pseudoautosomal region, limiting recombination to prevent deleterious exchanges.^[71] Such reductions facilitate the accumulation of sex-specific genes on differentiated chromosomes, contributing to sexual dimorphism without compromising autosomal synapsis.^[72] Experimental knockouts of SC components reveal species-specific impacts on viability and fertility. In C. elegans, mutations in syp-3 disrupt SC assembly but yield viable animals with sterility due to chromosome missegregation.^[73] Conversely, in mice, Sycp2 knockouts cause complete male sterility from meiotic arrest, while females remain subfertile, indicating partial compensation by other axial elements.^[74] In budding yeast (Saccharomyces cerevisiae), zip1 mutants lacking SC are viable with reduced spore viability, demonstrating that SC absence impairs but does not abolish recombination fidelity across taxa.^[75]

Pathological and Clinical Relevance

Implications in Cancer

Disruptions in the synaptonemal complex (SC) during meiosis can lead to faulty chromosome pairing and synapsis, resulting in nondisjunction of homologous chromosomes in germ cells and the production of aneuploid gametes.^[76] This meiotic error introduces genomic instability that may be inherited by offspring.^[77] In somatic cells, aberrant expression of SC proteins parallels meiotic functions and contributes to oncogenic processes. SYCP1, a core SC component, is frequently overexpressed in various cancers, including testicular germ cell tumors, where it forms SC-like structures that disrupt mitotic fidelity and promote aneuploidy.^[78] Similarly, SYCP3 mutations or dysregulation in non-small cell lung cancer enhance chromosomal instability by impairing homologous recombination repair, leading to higher rates of genomic rearrangements that drive tumor progression.^[79] These somatic re-expressions of meiotic machinery underscore how SC components can hijack normal DNA repair pathways to foster cancer hallmarks like uncontrolled proliferation.^[80] Therapeutically, targeting SC regulators offers potential for exploiting these vulnerabilities in cancer cells that mimic meiotic pathways. For example, ATM kinase, which coordinates SC assembly and double-strand break repair during meiosis, is often dysregulated in tumors; inhibiting ATM disrupts genome stability in cancer cells reliant on SC-like mechanisms, sensitizing them to DNA-damaging therapies.^[81] This finding positions SC components as biomarkers for aggressive disease and potential targets for metastasis-preventive interventions.^[82]

Associations with Reproductive Disorders

Defects in the synaptonemal complex (SC) are strongly associated with various reproductive disorders, primarily through disruptions in meiotic chromosome synapsis, recombination, and progression, leading to gamete production failure and infertility. In males, mutations in SC component genes such as SYCP3 have been linked to non-obstructive azoospermia, characterized by the absence of sperm in semen due to meiotic arrest. A heterozygous frameshift mutation (c.643delA) in SYCP3, resulting in a premature stop codon and C-terminal truncation, was identified in patients with azoospermia, causing impaired SC fiber formation and asynapsis that triggers spermatocyte apoptosis.^[83] In mouse models, complete knockout of Sycp3 similarly produces azoospermia via early meiotic arrest and spermatocyte loss, underscoring the conserved role of SYCP3 in SC integrity. In females, SC defects contribute to premature ovarian insufficiency (POI), defined as ovarian function loss before age 40, often involving accelerated follicle depletion and infertility. Variants affecting SC central element proteins, such as homozygous missense mutations in SYCE1 (e.g., c.613C>T), disrupt SC assembly and stability, leading to defective disassembly after pachytene and meiotic arrest in oocytes.^[84] Mouse models with Sycp1 knockout demonstrate analogous disassembly failures, with persistent SC fragments causing unrepaired DNA double-strand breaks, crossover defects, and complete infertility due to oocyte apoptosis. Although direct human SYCP1 variants are rare, SC gene disruptions collectively impair transverse filament function, exacerbating POI risk.^[85] Clinical diagnosis of SC-related infertility often involves testicular biopsies in males, where electron microscopy or spreading techniques reveal SC abnormalities such as fragmentation, asynapsis, or incomplete pairing in pachytene spermatocytes, correlating with maturation arrest and azoospermia or oligozoospermia. These findings distinguish SC defects from other spermatogenic failures and guide genetic counseling. In animal models, Sycp1-/- mice exhibit complete meiotic arrest at zygotene/pachytene, with no progression to metaphase I and total sterility in both sexes, highlighting the essentiality of SYCP1 for SC-mediated synapsis and recombination. SC mispairing has been implicated in genetic syndromes like 46,XX testicular disorder of sex development (DSD), where individuals with an XX karyotype develop male phenotypes but often face infertility due to sex chromosome pairing failures during meiosis. In such cases, SC analysis shows unpaired or mispaired autosomes and sex chromosomes, contributing to germ cell loss and azoospermia.^[86] Recent advances, including a 2023 genome-wide association study (GWAS), have identified variants in SC genes like SYCE2 that influence meiotic recombination rates and associate with idiopathic infertility and pregnancy loss, providing new genetic markers for unexplained reproductive failures.^[87] A 2025 study further identified a novel loss-of-function variant in SYCP2 associated with autosomal dominant male infertility.^[88]

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Aug 23, 2019 · Our data show that SYCP3 and the SYCP2 C terminus, which are known to form filaments in vitro, form a compact core around which cohesin ...
[17]
Exo1 recruits Cdc5 polo kinase to MutLγ to ensure efficient meiotic ...
The Mlh1–Mlh3 (MutLγ) endonuclease complex is critical for crossover resolution ... Double-strand break resection and resolution of double holliday junctions.
[18]
Protecting double Holliday junctions ensures crossing over during ...
Sep 24, 2025 · Imperatively, the biased resolution of double Holliday junction (dHJ) intermediates into crossovers occurs within the synaptonemal complex (SC), ...
[19]
EXO1 promotes the meiotic MLH1-MLH3 endonuclease ... - Nature
May 3, 2025 · The breaks are repaired by resection, followed by strand invasion into a matched DNA molecule, which in meiosis is often the homologous ...
[20]
ATR is required to complete meiotic recombination in mice - Nature
Jul 5, 2018 · Here we report that the DNA damage response protein kinase ATR is crucial for meiotic recombination and completion of meiotic prophase in mice.
[21]
ATM signaling modulates cohesin behavior in meiotic prophase and ...
Mar 6, 2023 · ... ATM/ATR kinase phosphorylation site regions in DNA-damage-response proteins. ... recombination and affects synaptonemal complex ...<|separator|>
[22]
Mouse HORMAD1 and HORMAD2, Two Conserved Meiotic ...
Oct 23, 2009 · HORMAD1/2 staining is intense on unsynapsed chromosome axes but is much reduced on synapsed regions marked by SYCP1 (Figures 2A, 3A, 4 and 5).
[23]
Meiotic homologous chromosome alignment and its surveillance are ...
Finally, HORMAD1 plays a key role in the male mid-pachytene checkpoint and the female meiotic prophase checkpoint, both of which eliminate meiocytes with ...
[24]
Functions and Regulation of Meiotic HORMA-Domain Proteins - PMC
Apr 27, 2022 · In mammals, HORMAD2 is not required to promote DSB formation or synapsis, unlike HORMAD1, but is required to recruit the ATR kinase to ...
[25]
Structural analysis of the human SYCE2–TEX12 complex provides ...
Structural analysis of the human SYCE2–TEX12 complex provides molecular insights into synaptonemal complex assembly ... Meiosis in mice without a synaptonemal ...
[26]
A Novel Mouse Synaptonemal Complex Protein Is Essential for ...
May 26, 2011 · The synaptonemal complex (SC) is a proteinaceous, meiosis ... Detailed analyses of Syce2 and Tex12 knockout mice showed that loss ...
[27]
https://doi.org/10.1038/nrg3573
[28]
https://doi.org/10.7554/eLife.21455
[29]
https://pmc.ncbi.nlm.nih.gov/articles/PMC9859615/
[30]
https://doi.org/10.1073/pnas.0406212102
[31]
https://doi.org/10.1146/annurev-genom-110122-090239
[32]
https://pmc.ncbi.nlm.nih.gov/articles/PMC8577265/
[33]
The organization, regulation, and biological functions of the ...
The synaptonemal complex (SC) is a meiosis-specific proteinaceous macromolecular structure that assembles between paired homologous chromosomes during meiosis ...
[34]
Kinetic analysis of synaptonemal complex dynamics during meiosis ...
Here, we directly measure the kinetics of Zip1-GFP assembly and disassembly in live cells of the yeast S. cerevisiae.
[35]
Synaptonemal Complex Components Persist at Centromeres and ...
Jun 28, 2012 · We show here that the SYCP1 and SYCP3 proteins are required for centromere pairing in mouse. Our findings define structures and possible mechanisms.Missing: stability | Show results with:stability
[36]
The APC/C activator FZR1 is essential for meiotic prophase I in mice
Fizzy-related 1 (FZR1) is an activator of the Anaphase promoting complex/cyclosome (APC/C) and an important regulator of the mitotic cell division cycle.Results · Fzr1 Loss Leads To A Small... · Role Of Apc/c During Meiotic...<|separator|>
[37]
Cyclic AMP in oocytes controls meiotic prophase I and primordial ...
Inhibiting cAMP synthesis in fetal ovaries delayed oocyte meiotic progression and inhibited the disassembly and degradation of synaptonemal complex protein 1.
[38]
Meiotic Chromosome Synapsis in Yeast Can Occur Without Spo11 ...
Previous reports have shown that the synaptic structures, the synaptonemal complexes (SCs), do not occur in yeast cells with the SPO11 gene removed. The Spo11 ...Missing: persistent | Show results with:persistent
[39]
The telomere bouquet facilitates meiotic prophase progression and ...
Nov 7, 2017 · At the chromosome oscillation stage, meiotic recombination and synaptonemal complex formation between homologous chromosomes are initiated.
[40]
Chromatin binding by HORMAD proteins regulates meiotic recombination initiation | The EMBO Journal
Summary of each segment:
[41]
Building the synaptonemal complex: Molecular interactions
Jul 20, 2023 · Here, we review what is known about CR-axis interactions in meiotic model organisms, with a focus on insight from C. elegans.<|control11|><|separator|>
[42]
https://genesdev.cshlp.org/content/34/23-24/1562.long
[43]
A new role for the synaptonemal complex in the regulation of meiotic ...
The synaptonemal complex (SC) is a meiosis-specific structure formed between homologous chromosomes during prophase that promotes DSB formation and biases ...
[44]
A new role for the synaptonemal complex in the regulation of meiotic ...
Dec 1, 2020 · The synaptonemal complex (SC) is a meiosis-specific structure formed between homologous chromosomes during prophase that promotes DSB formation and biases ...
[45]
Distribution of crossing over on mouse synaptonemal ... - PubMed
MLH1 is a mismatch repair protein necessary for meiotic recombination in mice, and MLH1 foci have been proposed to mark crossover sites. We present evidence ...
[46]
The synaptonemal complex imposes crossover interference and ...
We show that the synaptonemal complex, a structure that zips homologous chromosomes together during meiosis, is essential for crossover interference in ...<|control11|><|separator|>
[47]
Crossover/Noncrossover Differentiation, Synaptonemal Complex ...
Yeast mutants lacking meiotic proteins Zip1, Zip2, Zip3, Mer3, and/or Msh5 (ZMMs) were analyzed for recombination, synaptonemal complex (SC), and meiotic ...
[48]
Crossover assurance and crossover interference are ... - PubMed
Crossover assurance and crossover interference are distinctly regulated by the ZMM proteins during yeast meiosis ... synaptonemal complex formation. Unlike ...
[49]
The choice in meiosis – defining the factors that influence crossover ...
Feb 15, 2011 · Crossover assurance: The mechanism that makes certain each homologous chromosome pair will receive at least one meiotic crossover. Crossover ...
[50]
Crossover Patterning by the Beam-Film Model - Research journals
We previously-described a mechanical “stress and stress relief” model for CO patterning with an accompanying mathematical description (the “beam-film model”).
[51]
Meiosis in Mice without a Synaptonemal Complex | PLOS One
... SYCP3 double-null oocytes contained 1–3 chiasmata per oocyte. This corresponds to the number of chiasmata observed in MLH1- and MLH3-null oocytes [40], [41].<|control11|><|separator|>
[52]
Meiotic interference among MLH1 foci requires neither an intact ...
Mar 1, 2007 · Crossovers display interference, i.e. their spacing is more even than would be expected if they were placed randomly along the chromosomes.
[53]
Conservation and Variability of Meiosis Across the Eukaryotes
Nov 23, 2016 · However, it is possible that the synaptonemal complex and the delicate crossover control related to its presence were later acquisitions. Later ...
[54]
Unconventional conservation reveals structure-function ... - eLife
Nov 17, 2021 · This article overcomes this challenge by developing and validating a clever protein structure-based approach that leverages sequence divergence.Missing: sensor | Show results with:sensor
[55]
From conservation to adaptation: understanding the synaptonemal ...
Following its first description in 1958 by Montrose J. Moses in the salamander Plethodon cinereus [19], the SC has been described in hundreds of species. Due to ...Missing: discovery | Show results with:discovery
[56]
Hydra meiosis reveals unexpected conservation of structural ... - PNAS
Sep 24, 2012 · The synaptonemal complex (SC) is a key structure of meiosis, mediating the stable pairing (synapsis) of homologous chromosomes during ...
[57]
Holding it together: rapid evolution and positive selection in the ...
May 5, 2016 · The synaptonemal complex (SC) is a highly conserved meiotic structure that functions to pair homologs and facilitate meiotic recombination ...
[58]
Conservation and Variability of Synaptonemal Complex Proteins in ...
Hydra meiosis reveals unexpected conservation of structural synaptonemal complex proteins across metazoans. Proceedings of the National Academy of Sciences ...
[59]
The synaptonemal complex central element SCEP3 interlinks ...
Jun 27, 2025 · Regulation of the MLH1–MLH3 endonuclease in meiosis. ... SYCE2 is required for synaptonemal complex assembly, double strand break repair, and ...
[60]
Mouse Sycp1 functions in synaptonemal complex assembly, meiotic ...
We disrupted the mouse gene encoding TF protein Sycp1 to analyze the role of TFs in meiotic chromosome behavior and recombination. Sycp1-/- mice are infertile, ...Results · Early Meiosis In Sycp1... · Role Of Sycp1 In Later Steps...
[61]
Sex in Symbiodiniaceae dinoflagellates: genomic evidence ... - Nature
Jun 17, 2020 · We report the absence of genes that encode main components of the synaptonemal complex (SC), a protein structure that mediates homologous chromosomal pairing ...
[62]
Meiosis-associated expression patterns during starvation-induced ...
May 22, 2025 · ZIP1 and RED1 are both involved in the synaptonemal complex, which is not strictly needed for meiosis in many taxa [43]. The common absence ...
[63]
The Dynamics of Homologous Chromosome Pairing during Male ...
In male Drosophila melanogaster, meiosis occurs in the absence of recombination or a recognizable synaptonemal complex (SC).
[64]
MNM and SNM maintain but do not establish achiasmate homolog ...
May 28, 2019 · In contrast to canonical meiosis, homologous chromosome pairing does not include the formation of a synaptonemal complex and of cross-overs in ...
[65]
Achiasmy: Male Fruit Flies Are Not Ready to Mix - PMC
Jul 19, 2016 · Since there is no recombination during Drosophila male meiosis the synaptonemal complex is undetectable. However, the pairing sites on the ...
[66]
The synaptonemal complex has liquid crystalline properties and ...
Jan 3, 2017 · The synaptonemal complex (SC) is a polymer that spans ~100 nm between paired homologous chromosomes during meiosis.Missing: width | Show results with:width
[67]
The Synaptonemal Complex Shapes the Crossover Landscape ...
The SC has been demonstrated to have at least two conserved functions in meiotic prophase. First, the SC serves to stabilize and maintain tight associations ...Missing: review | Show results with:review
[68]
Meiosis in Polyploids and Implications for Genetic Mapping: A Review
Variations in meiosis are mainly due to the type of presynaptic alignment, either distal or complete; the number and distribution of synapsis initiation points; ...7. Meiosis In Allopolyploids · 13. The Zip4 Gene · 18. Meiotic Proteins And...
[69]
The synaptonemal complex stabilizes meiosis in allotetraploid ...
We studied the role of the synaptonemal complex in polyploid meiosis by mutating the transverse filament component ZYP1 in allotetraploid Brassica napus and ...
[70]
A search for synaptonemal complexes in Ustilago maydis
Aug 1, 1981 · It seems that the SC is not necessary for recombination repair of DNA broken by irradiation. DISCUSSION. Dr P. B. Moens (personal ...
[71]
Bioinformatic identification of Ustilago maydis meiosis genes
Aug 7, 2025 · We report the absence of genes that encode main components of the synaptonemal complex (SC), a protein structure that mediates homologous ...
[72]
The homologous recombination system of Ustilago maydis - PMC
In many systems meiotic cells develop specialized higher order chromosome structures culminating in formation of the synaptonemal complex (SC), the tripartite ...
[73]
The Chromosomes of Birds during Meiosis - Karger Publishers
Dec 29, 2016 · This review highlights the main contributions of synaptonemal complex and lampbrush chromosome research to the current knowledge of avian ...Crossover Markers Along... · The Effects Of Abnormal... · The Zw Pair During Meiosis
[74]
Heteromorphic sex chromosomes: Navigating meiosis without a ...
Jul 27, 2011 · During synapsis, the synaptonemal complex (green cross lines) stabilizes the aligned homologous chromosomes. In this context, crossover ...
[75]
SYP-3 Restricts Synaptonemal Complex Assembly to Bridge Paired ...
The syp-3(ok758) deletion allele was generated by the C. elegans Gene Knockout Consortium. In parallel, F39H2.4 was identified through a functional genomics ...Missing: viable sterile<|separator|>
[76]
Mouse SYCP2 is required for synaptonemal complex assembly and ...
The fertility of homozygous Sycp2 mutant mice is sexually dimorphic; males are sterile because of a block in meiosis, whereas females are subfertile with ...Mouse Sycp2 Is Required For... · Results · Sycp2 Is Associated With...
[77]
Germ Cell Differentiation and Synaptonemal Complex Formation ...
Breedings of heterozygous mice pro- duced offspring in the expected Mendelian distribution are under CPEB-mediated translational control. Page 3. CPEB Knockout ...
[78]
Disruption of pairing and synapsis of chromosomes causes stage ...
Aug 6, 2025 · Incorrect assembly of the synaptonemal complex results in cell death, impaired meiotic recombination and aneuploidy. Oocytes with meiotic ...
[79]
Chromosomal Instability in Genome Evolution: From Cancer to ...
Feb 7, 2023 · Genomic instability in germ cells can lead to chromosomal anomalies, resulting in sterility or genetic diseases in the offspring [3,4,5], but ...2.4. Complex Genomic... · 3.3. Chromosomal Variance In... · 6. Germ Cells And Meiotic...<|control11|><|separator|>
[80]
Cancer Risk in Children with Birth Defects and in Their Families
We observed an increased overall cancer risk in individuals with birth defects. The highest risks were seen for individuals with malformations in the nervous ...Missing: meiotic | Show results with:meiotic
[81]
Centrosome dysfunction associated with somatic expression of the ...
Dec 8, 2021 · Here we report that meiotic synaptonemal complex protein TEX12 is frequently mis-expressed in cancer. We also show that TEX12 has a ...
[82]
Synaptonemal complex protein 3 is associated with ...
Jun 17, 2017 · Synaptonemal complex protein 3 is associated with lymphangiogenesis in non-small cell lung cancer patients with lymph node metastasis.
[83]
Synaptonemal complex proteins modulate the level of genome ...
Dec 31, 2020 · Accumulating evidence indicates that they are epigenetically expressed in cancers of various origin. Not surprisingly, in contrast to their ...
[84]
ATM/ATR kinases link the synaptonemal complex and DNA double ...
ATM/ATR kinases link the synaptonemal complex and DNA double-strand break repair pathway choice - PMC.
[85]
Genetic predispositions to childhood leukemia - PMC - NIH
This review focuses on describing the molecular basis of multiple germline syndromes that confer an increased risk of leukemia, and will specifically highlight ...
[86]
Immunological modifications & delayed ovarian cancer recurrence
Jun 25, 2023 · We report for the first time, the association between immunological modifications in response to chemotherapy and the time of recurrence.
[87]
Meiotic protein SYCP2 confers resistance to DNA-damaging agents ...
Feb 21, 2024 · Here, we show that SYCP2, a meiotic protein in the synaptonemal complex, is aberrantly and commonly expressed in breast and ovarian cancers and ...
[88]
Azoospermia in patients heterozygous for a mutation in SYCP3
Sycp3 encodes a component of the synaptonemal complex. A null mutation of Sycp3 in mice causes azoospermia with meiotic arrest. We tested the hypothesis ...
[89]
Exome Sequencing Reveals SYCE1 Mutation Associated With ...
These results highlight the importance of the synaptonemal complex and meiosis in ovarian function. Issue Section: JCEM Online: Advances in Genetics. Collection ...
[90]
Alterations in synaptonemal complex coding genes and human ...
Feb 21, 2022 · SYCP2 and SYCP3 are the core axis proteins in mammals and have a direct interaction [26, 67]. SYCP2 has 47 exons and encodes a 1350 amino acid ...<|separator|>
[91]
Diseases associated with SYCP1 | Open Targets Platform
synaptonemal complex protein 1. Ensembl: ENSG00000198765UniProt: Q15431GeneCards: SYCP1HGNC: SYCP1Project ... 46,XX testicular disorder of sex development.
[92]
Variant in the synaptonemal complex protein SYCE2 associates with ...
Jan 29, 2024 · SYCE2 is part of the synaptonemal complex, a protein structure that mediates alignment, synapsis (pairing) and recombination of homologous ...