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Pronucleus

A pronucleus is the haploid derived from either the or following fertilization, prior to the fusion of their genetic material to form the diploid . In this stage, the male pronucleus forms when the 's highly condensed decondenses after entering the , while the female pronucleus arises from the 's after completion of II. Each pronucleus contains 23 chromosomes and is enclosed by a , representing a critical in the restoration of diploidy during reproduction. The formation of pronuclei is triggered by specific biochemical changes post-fertilization. In sea urchins, the chromatin decondenses through the of sperm-specific histones. In mammals, decondensation is facilitated by factors from the egg such as , which reduces bonds in protamines, followed by the replacement of protamines with histones from the , allowing the male pronucleus to expand and mature over several hours. The female pronucleus, meanwhile, forms as the extrudes the second , ensuring its haploid state. These transformations prepare the genetic material for syngamy, the union of paternal and maternal genomes. Pronuclear migration involves cytoskeletal dynamics that bring the two nuclei into close proximity. In sea urchins, the sperm organizes into an that contacts and pulls the pronucleus toward the male, with completing in under an hour. Mammalian zygotes exhibit a slower process, lasting about 12 hours, driven by astral from the and actin-based movements, during which often begins within the pronuclei. This ensures proper alignment for genetic fusion and is essential for embryonic development; disruptions can lead to or developmental arrest. The fusion of pronuclei, or syngamy, culminates in the formation of the . In some like sea urchins, the pronuclear envelopes break down, allowing direct merging of the haploid nuclei into a single diploid . In mammals, including humans, the pronuclei approach each other but do not fully fuse; instead, their envelopes interdigitate, and the chromosomes condense and align on a shared mitotic during the first division, resulting in diploid daughter cells. This process typically occurs in the ampullary region of the in humans. Abnormalities in pronuclear formation or migration, such as asynchronous development, are associated with and are key indicators in assisted reproductive technologies like fertilization.

Definition and Characteristics

General Definition

A pronucleus is the haploid nucleus derived from either the or during fertilization, containing a single set of chromosomes that prepares the parental genetic material for fusion into the diploid nucleus. In humans, each pronucleus carries 23 chromosomes, half the diploid number found in cells. Upon fusion during syngamy, these combine to form the zygote nucleus with 46 chromosomes, marking the initiation of embryonic . Pronuclei differ from the nuclei of mature gametes, which are produced through meiosis and remain in a condensed state until fertilization. These structures are transient, existing only briefly to facilitate the reorganization and alignment of genetic material prior to nuclear fusion. The formation and function of pronuclei represent a conserved feature of sexual reproduction across eukaryotes, occurring in diverse taxa such as animals, plants, and algae as an essential prelude to syngamy.

Structural Features

The pronucleus constitutes an enlarged haploid nucleus enveloped by a reformed nuclear membrane, enclosing decondensed chromatin organized into a less compact structure compared to the preceding gametic nuclei. This nuclear envelope, derived from oocyte endoplasmic reticulum components, integrates proteins such as lamins to provide structural support and facilitate nucleocytoplasmic transport, while the chromatin incorporates maternal histones and associated epigenetic modifiers. Within the pronucleus, nucleoli or nucleolar precursor bodies emerge as prominent substructures, clustering near the chromatin and containing ribosomal RNA precursors essential for early embryonic transcription. In mammals, the male pronucleus typically measures approximately 20-25 μm in diameter and arises from the sperm head, where protamine-packaged DNA undergoes rapid remodeling to histone-based chromatin supplied by the oocyte cytoplasm. This structure initially lacks embedded centrioles, which are contributed by the sperm but remain in the adjacent cytoplasm until post-fusion integration during syngamy. The male pronucleus often exhibits distinct epigenetic marks, such as elevated histone acetylation and reduced methylation on H3K9, contributing to its unique chromatin accessibility. The female pronucleus, by contrast, is similarly sized at around 20-30 μm in diameter in species like humans and mice, but features more prominent and numerous nucleoli due to maternal accumulation of ribosomal components. It harbors substantial maternal stores of RNAs and proteins, including factors like nucleoplasmin that aid in chromatin organization, and may appear asymmetrically shaped owing to its proximity to the oocyte's polar bodies and cytoplasmic determinants. This asymmetry influences nucleolar distribution and overall pronuclear morphology without affecting its core haploid composition. Pronuclei are readily observable in living mammalian zygotes through phase-contrast or differential interference contrast light microscopy, revealing their spherical to ovoid contours and internal nucleolar spots. For detailed visualization, fluorescent staining with DNA-binding dyes such as highlights decondensed patterns and nucleolar exclusion zones under epifluorescence or , enabling non-invasive assessment of structural integrity.

Formation

Female Pronucleus Formation

The formation of the female pronucleus is initiated by the entry of the into the , which triggers a series of signaling events including oscillatory increases in intracellular calcium levels that propagate as maternal calcium waves. These calcium signals induce cortical granule , a process that releases enzymes into the perivitelline space to modify the and prevent by hardening the egg coat. Concurrently, the calcium oscillations resume and drive the completion of the second meiotic division in the oocyte, which had been arrested at II. Upon completion of meiosis II, the oocyte extrudes the second polar body, retaining a haploid set of chromosomes in the ooplasm that subsequently decondense to form the female pronucleus. This decondensation involves the remodeling of the highly condensed oocyte chromatin, facilitated by maternal factors that promote histone modifications and chromatin relaxation, allowing the assembly of a functional nuclear envelope around the haploid genome. In mammals, this process is supported by the incorporation of maternal histone chaperones and variants, which ensure proper chromatin structure without the extensive protamine-to-histone exchange required in the male counterpart. The timeline for female pronucleus formation in mammals typically occurs within 3 to 10 hours post-fertilization, with species-specific variations; for instance, in mice, it forms around 7.5 hours after , while in humans, it appears by approximately 8 hours on average. This developmental window involves reorganization to facilitate extrusion and pronuclear assembly, coordinated by the calcium waves that sustain egg activation. Across species, the process exhibits notable variations. In sea urchins, where the oocyte completes meiosis prior to fertilization, the female pronucleus forms rapidly, within less than 30 minutes post-sperm entry, due to the pre-existing haploid state and swift decondensation.

Male Pronucleus Formation

Following sperm-egg fusion, the acrosome-reacted sperm penetrates the zona pellucida and delivers its haploid nucleus into the ooplasm, where initial structural changes occur to prepare for pronuclear development. The acrosome reaction, triggered by zona proteins, exposes enzymes that facilitate zona traversal and enables plasma membrane fusion with the egg. Upon entry, disassembly of the perinuclear theca—a cytoskeletal layer encapsulating the sperm nucleus—exposes the protamine-packaged DNA, marking the onset of nuclear remodeling. Decondensation of the sperm follows, involving the exchange of protamines for -derived s to restore a nucleosomal structure compatible with transcription. This process is driven by oocyte factors, notably , which reduces bonds in protamines, allowing chromatin uncoiling and histone incorporation. A new then assembles around the decondensed DNA, incorporating s for transport. Key proteins such as importins facilitate nuclear pore complex by regulating nucleoporin , while SMC proteins (structural maintenance of chromosomes) contribute to chromatin looping and higher-order during this reformation. Concurrently, the sperm's proximal serves as the primary microtubule-organizing center, nucleating astral essential for subsequent zygotic events. In humans, male pronucleus formation is typically observed between 8-12 hours post-sperm entry, as seen in (ICSI) procedures where decondensation begins around 4 hours in the . This timeline varies across species; in mammals, oocyte is crucial for and efficient decondensation, whereas in like sea urchins, the process is faster—often under 1 hour—due to the acidic cytoplasmic environment that promotes dissociation and .

Role in Fertilization

Pronuclear Migration

Pronuclear migration refers to the cytoskeletal-driven process in which the male and female pronuclei move toward each other within the following fertilization, positioning them for subsequent syngamy. This movement is essential for the spatial alignment of parental genomes and is mediated primarily by networks organized by the sperm-derived , which forms a radial to generate pulling forces on the pronuclei. motors, anchored to the cortex or organelles, walk along these to produce the necessary force for pronuclear translocation, often in coordination with polymerization and depolymerization dynamics. In mammalian zygotes, such as those of mice, pronuclear migration occurs in two distinct phases: a rapid peripheral driven by polymerization via Formin-2 and nucleators, which propels the male pronucleus inward from the fertilization cone, followed by a slower central reliant on microtubule-dynein interactions for . The sperm-introduced proximal duplicates to form acentriolar microtubule-organizing centers (aMTOCs) that nucleate the , facilitating capture and transport of both pronuclei without a strict centriole dependency in all cases. This process typically unfolds over 4-8 hours post-fertilization in mice, with the male pronucleus starting farther from the cell center (approximately 37 μm) and migrating at initial velocities of 0.38 μm/min, slowing to 0.01 μm/min as it approaches the female pronucleus. Calcium signaling plays a regulatory role in pronuclear migration across species, with oscillatory calcium waves—often triggered by inositol 1,4,5-trisphosphate (IP3) receptors—promoting microtubule polymerization and aster formation shortly after fertilization. In mammalian eggs, these calcium oscillations, initiated by sperm phospholipase C zeta, sustain the cytoskeletal rearrangements needed for migration, while in sea urchins, a secondary calcium wave originating at the sperm entry point coincides with the onset of pronuclear movement. Species-specific variations highlight diverse cytoskeletal adaptations; in sea urchins, migration is microtubule-based via a rapidly maturing sperm aster, completing in under 30 minutes with velocities up to 4.9 μm/min, driven by dynein-mediated pulling without pronounced involvement in the central phase. In contrast, like Pelvetia, the female pronucleus remains anchored near the cell center by a stable and F- network, while the sperm pronucleus migrates toward it along fixed tracks, emphasizing a more static, directed pathway compared to the dynamic in mammals. Time-lapse imaging techniques, including 3D with fluorescent labels for (e.g., H2B-mCherry) and cell membranes (e.g., MyrGFP), have revealed synchronous rotation and approximation of pronuclei during , with the pronucleus often rotating as it is pulled centrally in mammalian zygotes. These visualizations demonstrate coordinated oscillatory movements and highlight the precision of dynein-driven transport in aligning the pronuclei.

Syngamy and Genome Activation

Syngamy represents the culminating event of fertilization, wherein the pronuclei approximate each other, their nuclear envelopes break down, and the parental align on a shared mitotic to achieve diploidy in the daughter cells of the first cleavage division. This process is triggered by the activation of (MPF), which induces condensation and the assembly of a mitotic . In mammalian zygotes, the formation often involves dual structures that initially align maternal and paternal separately before converging, ensuring proper during the impending first mitotic division. The timeline of syngamy in humans typically occurs approximately 20 to 24 hours post-insemination, coinciding with the completion of in the pronuclei. This is preceded by the disassembly of the pronuclear lamina, a meshwork of proteins that supports the , facilitating envelope breakdown and release into the shared . Prior to this, initiates independently within each pronucleus during the S-phase, a process observed in species such as mice and humans, where replication licensing and origin firing occur asynchronously between parental genomes. Additionally, karyopherin-mediated nuclear import, which facilitates protein entry into the pronuclei during their formation, halts as the envelopes disassemble, marking the transition to mitotic events. Concomitant with syngamy is the onset of zygotic genome activation (ZGA), part of the broader maternal-to-zygotic transition (MZT) that shifts developmental control from maternal transcripts to embryonic . Recent research as of 2025 indicates that ZGA initiates at the one-cell stage in both mice and humans, with low-level transcription detectable shortly after fertilization, followed by a minor wave during the pronuclear stage that involves activity primarily from the male pronucleus in mice. The major wave follows at the 2-cell stage in mice and 4- to 8-cell stage in humans. The successful alignment and processing during syngamy terminates the pronuclear phase and initiates embryonic cleavage, with the integrated now poised for robust transcriptional activation.

Historical Development

Early Discoveries

The discovery of the pronucleus built upon foundational 19th-century studies of gametes, notably von Baer's 1827 identification of the mammalian ovum through microscopic examination of ovarian follicles in dogs and other mammals. This breakthrough shifted embryological research toward understanding egg structure and fertilization processes in animals. In 1875, Belgian cytologist Edouard Van Beneden advanced this field by observing pronuclei in the oocytes of rabbits and bats using light microscopy on fixed specimens. His work, detailed in a paper on egg maturation and early embryonic development, described the formation of a peripheral male pronucleus from sperm material diffusing through the egg membrane and a central female pronucleus, which subsequently fused to form the first embryonic nucleus. Van Beneden's observations linked these pronuclei to the broader context of egg maturation, foreshadowing connections to meiotic processes observed in his later studies. One year later, in 1876, German zoologist Oscar Hertwig provided the first clear visualization of pronuclear fusion during fertilization in echinoderms through experiments on eggs. By combining and eggs in seawater and observing the process under a —exploiting the transparency of eggs—Hertwig documented the pronucleus entering the egg and fusing with the female pronucleus, resolving debates on the role of in . These early efforts relied on rudimentary techniques such as direct microscopic viewing of living or freshly dissected material, basic fixation methods, and emerging nuclear staining to enhance visibility, without the benefit of molecular tools. Collectively, Van Beneden's and Hertwig's discoveries established pronuclei as distinct entities central to fertilization, transforming the view of the process from mere union to a precise merger essential for embryonic .

Key Milestones

In the and , electron microscopy provided the first detailed views of pronuclear , revealing the decondensation of the nucleus into a fibrous network within the , as observed in mammalian species like rabbits. Pioneering studies by C.R. Austin in 1961 described these processes, highlighting the transformation of the compact head into a swollen pronucleus through and reformation. During the 1970s, research on advanced models of , where two sperm nuclei migrate to distinct targets in the embryo sac, forming separate pronuclei that fuse to initiate endosperm development in angiosperms. These cross-species investigations, using species like and lilies, elucidated pronuclear patterns and mechanisms, contrasting with animal systems and informing evolutionary comparisons. The 1980s marked the IVF era, where pronuclei served as visible markers of successful fertilization in embryos, with early observations documenting their formation and symmetry within 18-24 hours post-insemination. Studies in by teams including Trounson and colleagues reported the timing of pronuclear appearance in cultured zygotes, enabling non-invasive assessment of fertilization viability. In the 1990s and 2000s, molecular insights deepened, with 1993 experiments on the alga Pelvetia demonstrating waves that trigger pronuclear formation and migration during fertilization. Concurrently, research in the 2000s uncovered the mechanisms of removal from the male pronucleus, linking it to epigenetic reprogramming via glutathione-mediated disulfide bond reduction and reassembly. Post-2010 advancements included applications, where pronuclear injections into enabled targeted to study paternal and maternal contributions, as shown in 2015 human tripronuclear experiments achieving high-efficiency mutations. Live-cell imaging techniques in 2015 further revealed the role of mammalian centrioles in pronuclear migration, illustrating microtubule-dependent centering of the male pronucleus toward the female.

Clinical and Research Significance

In Assisted Reproduction

In assisted reproduction, particularly fertilization (IVF), the observation of pronuclei serves as a critical indicator of successful fertilization. Approximately 16-18 hours after , embryologists assess for the presence of two distinct pronuclei (2PN), which signifies normal monospermic fertilization where one has penetrated the oocyte, leading to the decondensation of the head into the pronucleus and the formation of the pronucleus. The appearance of exactly two pronuclei is considered the ideal marker for selecting embryos suitable for transfer or further culture, as it confirms the extrusion of the second and the initiation of syngamy. In (ICSI) procedures, this assessment similarly verifies successful injection, though ICSI typically exhibits lower rates of compared to conventional IVF. The evaluation of pronuclei extends beyond mere counting to include morphological characteristics such as size symmetry and the of nucleolar precursor bodies (NPBs), which provide predictive insights into viability and implantation potential. Zygotes with symmetrical pronuclei and evenly distributed, large NPBs (e.g., in a polarized ) are associated with higher developmental competence and improved implantation rates. A study evaluating pronuclear in conjunction with subsequent grading demonstrated that selecting based on these features significantly enhances implantation success. These assessments help prioritize embryos for transfer, reducing the risk of selecting non-viable ones in cycles where multiple oocytes are fertilized. Non-invasive techniques, such as time-lapse imaging systems integrated into incubators, have revolutionized pronuclei monitoring by allowing continuous observation without removing from optimal culture conditions. These systems capture images at intervals of 5-20 minutes, enabling the tracking of pronuclear formation, alignment, and disappearance in real-time, which correlates with subsequent timing and quality. By minimizing environmental disturbances, time-lapse monitoring improves the accuracy of fertilization confirmation and supports dynamic embryo selection algorithms that integrate pronuclear dynamics with later morphokinetic parameters. Recent advancements as of 2025 include AI-based analysis of pronuclear patterns to further refine viability predictions. Abnormal pronuclear configurations, such as the presence of three or more pronuclei (3PN), are indicative of , where multiple sperm enter the , and such zygotes are routinely discarded due to their high risk of chromosomal abnormalities and poor developmental outcomes. In human IVF , approximately 70-80% of mature oocytes achieve normal fertilization as evidenced by the formation of two pronuclei, reflecting standard laboratory benchmarks for successful rates. This statistic underscores the efficiency of modern protocols while highlighting the need for careful pronuclear evaluation to optimize cycle success.

Abnormalities and Implications

One common abnormality in pronucleus formation is , where more than one fertilizes the , resulting in multiple male pronuclei (typically three or more in IVF settings). This occurs due to failure of the , a key polyspermy-blocking mechanism involving of cortical granules that modifies the to prevent additional penetration. leads to triploid zygotes with an extra paternal set, causing embryonic arrest at early stages and inviable development. Asynchronous pronuclear development, characterized by delayed formation or differing sizes of the male and female , often stems from dysregulation of calcium oscillations essential for and pronuclear decondensation. Genetic defects in factors, such as in PLCZ1 encoding , can impair these oscillations, leading to incomplete and asynchrony. Such abnormalities reduce viability, increasing rates of implantation failure and early arrest. Structural anomalies in pronuclei, including fragile nuclear envelopes and uneven distribution, are frequently linked to defects like deficiency, which disrupt proper decondensation and reprogramming during pronuclear formation. These issues can precipitate epigenetic errors, such as aberrant at imprinted loci, contributing to imprinting disorders like Beckwith-Wiedemann or Silver-Russell syndromes in ART-conceived offspring. In research, pronucleus abnormalities serve as models for studying mechanisms, as multipronucleate zygotes often exhibit chromosomal imbalances that mimic early embryonic errors. Clinically, they elevate risk, accounting for a significant portion of IVF cycle failures due to non-viable embryos. These abnormalities are more prevalent than , where natural barriers limit sperm access; for instance, eggs employ a rapid electrical fast-block to prevent , a less emphasized in mammals but complemented by zona modifications.

References

  1. [1]
    Fusion of the Genetic Material - Developmental Biology - NCBI - NIH
    The egg nucleus, once it is haploid, is called the female pronucleus. Once inside the egg, the sperm nucleus decondenses to form the male pronucleus. The sperm ...Missing: definition | Show results with:definition
  2. [2]
    Embryology, Week 1 - StatPearls - NCBI Bookshelf
    Apr 17, 2023 · Once the sperm penetrates the ovum, the cytoplasm of the mature ovum contains two pronuclei: a male pronucleus composed of a sperm head with its ...
  3. [3]
    Fertilization - Molecular Biology of the Cell - NCBI Bookshelf - NIH
    The pronuclei migrate toward the center of the egg. When they come together, their nuclear envelopes interdigitate. The centrosome replicates, the nuclear ...
  4. [4]
    Glossary - Exploring the Biological Contributions to Human Health
    Haploid. The number of chromosomes in a sperm or egg cell, half the diploid number. In humans, the haploid number is 23. Haplotype. 1. A set of alleles of a ...
  5. [5]
    The Human Genome - NCBI - NIH
    These are called somatic cells, in contrast to sex cells or gametes, which are haploid and have just 23 chromosomes, comprising one of each autosome and one sex ...
  6. [6]
    Glossary - Heritable Human Genome Editing - NCBI Bookshelf - NIH
    Sep 3, 2020 · Pronucleus. The haploid nucleus of an oocyte or sperm, either prior to fertilization or immediately after fertilization, before the sperm and ...
  7. [7]
    Gamete Nuclear Migration in Animals and Plants - PMC
    Apr 24, 2019 · In animals, gamete nuclei during the process of fertilization are termed pronuclei. By contrast, the term pronucleus is hardly employed in ...
  8. [8]
    Fertilization Mechanisms in Flowering Plants - PMC - PubMed Central
    The sperm aster captures the female pronucleus and the two pronuclei rapidly move towards each other and to the center of the oocyte in a dynein-mediated ...
  9. [9]
    Nuclear envelope dynamics during male pronuclear development
    The poreless sperm nuclear envelope is replaced by a functional male pronuclear envelope and the highly compact male chromatin decondenses. ... decondensation ...
  10. [10]
    Potential of zygotes to produce live births can be identified by the ...
    Apr 12, 2017 · The average sizes of the female and male pronuclei 8 hours prior to the PNMBD were 356.3 μm2 (±48.3) and 404.8 μm2 (±61.2), respectively. They ...
  11. [11]
    Two mechanisms drive pronuclear migration in mouse zygotes
    Feb 5, 2021 · The male pronucleus assembles within the fertilization cone and is rapidly moved inwards by the flattening cone. Rab11a recruits the actin ...
  12. [12]
    Remodelling the paternal chromatin at fertilization in mammals - PMC
    After fertilization, the protamines are replaced by oocyte-supplied histones. This replacement occurs as the oocyte completes the second meiotic division and ...
  13. [13]
    Developmental Stages in Human Embryos
    Although "in most mammalian species, the male pronucleus has been reported to be larger than the female pronucleus," the converse has been found in one ...
  14. [14]
    Localisation of RNAs and proteins in nucleolar precursor bodies of ...
    Sep 17, 2015 · Early embryos of all mammalian species contain morphologically distinct but transcriptionally silent nucleoli called the nucleolar precursor bodies (NPBs).
  15. [15]
    Single nucleolus precursor body formation in the pronucleus of ...
    Aug 20, 2018 · Here, we show that in mouse embryos, the volume of NPB materials plays a major role in the NPB scaling through a limiting component mechanism ...
  16. [16]
    Visualization of pronuclei in living bovine zygotes - PubMed
    The one-celled ova were treated with 4'-6'-diamidino-2-phenylindole (DAPI) and observed under ultraviolet light by fluorescence microscopy. Both male and ...
  17. [17]
    Re-starting life: Fertilization and the transition from meiosis to mitosis
    Following fertilization, the egg exits from meiosis and assembles a haploid pronucleus. At the same time the sperm genome, which enters the egg in a highly ...Fertilization · Sperm-Egg Binding · Asymmetric Spindle...Missing: definition | Show results with:definition
  18. [18]
    Microtubule-Based Mechanisms of Pronuclear Positioning - PMC
    Feb 23, 2020 · Union of haploid male and female pronuclei in many animals occurs through rearrangements of the microtubule cytoskeleton into a radial array of microtubules ...Missing: definition | Show results with:definition
  19. [19]
    Pronucleus - an overview | ScienceDirect Topics
    Pronucleus refers to the stage of a fertilized egg where the sperm head has not yet merged with the egg nucleus, resulting in the formation of two distinct ...
  20. [20]
    A combination of maternal histone variants and chaperones ... - NIH
    Protamines are replaced with histones prior to zygotic genome activation and fusion of parental pronuclei. This exchange can be detected within a few minutes of ...
  21. [21]
    The timing of pronuclear formation, DNA synthesis and cleavage in ...
    Pronuclei formed between 3 and 10 h post-insemination (hpi; median 8 hpi). S-phase commenced between 8 and 14 hpi, and was completed between 10 and 18 hpi.
  22. [22]
    Central Cell in Flowering Plants: Specification, Signaling, and ...
    Oct 21, 2020 · Double fertilization entails fusion of sperm cells with the two dimorphic female gametes, the egg and the central cell. The potentiality of ...
  23. [23]
    Site of the mammalian sperm physiological acrosome reaction - NIH
    This reaction releases the acrosome's soluble lysins, which facilitate the creation of a path in the egg coat during penetration (although a variety of evidence ...
  24. [24]
    Defective sperm head decondensation undermines the success of ...
    Further, the transformation into a male PN requires among other things the swapping of the DNA-associated sperm's protamines with maternal histones as well as ...
  25. [25]
    Nucleosome assembly is required for nuclear pore complex ... - NIH
    In this study, we set out to reveal additional functions of nucleosome assembly and generated a ND paternal pronucleus in mouse zygotes through depletion of ...
  26. [26]
    Linker histone H1.8 inhibits chromatin binding of condensins and ...
    It has been shown that the sperm male pronucleus dynamically interacts ... SMC proteins and topo II. Materials and methods. Key resources table. Reagent ...
  27. [27]
    The Sperm Centrioles - PMC - PubMed Central - NIH
    E) The centrioles have a role in organizing the zygote's cytoskeleton - they form the astral microtubules (A-M) that mediate pronuclei migration and assist in ...
  28. [28]
    Kinetics of human male pronuclear development in a heterologous ...
    ... male pronucleus (mPN) within the egg cytoplasm. This process consists of complex overlapping stages [1–3]: (1) removal of the sperm nuclear envelope and ...
  29. [29]
    Calcium signals in and around the nucleus in sea urchin eggs
    In fact, a second calcium wave propagates through the egg as pronuclear migration begins; this wave also originates at the point of sperm entry.
  30. [30]
    Computer Simulations and Image Processing Reveal Length ...
    A male pronucleus migrates toward the center of an egg to reach the female pronucleus for zygote formation. This migration depends on microtubules growing ...
  31. [31]
    Principles of mammalian fertilization
    Maturation-promoting factor (MPF) must be activated to induce nuclear envelope breakdown, chromosome condensation and formation of the spindle apparatus.
  32. [32]
    Making sense out of syngamy at the onset of mammalian development
    Aug 2, 2018 · Dual-spindle formation in zygotes keeps parental genomes apart in early mammalian embryos. Science. 2018;361(6398):189–193. doi: 10.1126 ...
  33. [33]
    Syngamy, pronucleus, pronuclear breakdown and zygote
    About 5 h after insemination the PB2 is extruded and both PN enter the G1-phase (zygote stage). Thereafter, both PN enter the S-phase (DNA replication), ...Missing: timeline | Show results with:timeline
  34. [34]
    Nuclear envelope breakdown in mammalian cells involves stepwise ...
    Nuclear envelope breakdown involves stepwise lamina disassembly, starting with A-type lamins, and microtubule-driven deformation, with indentations and ...
  35. [35]
    Differential nuclear import sets the timing of protein access ... - Nature
    Oct 6, 2022 · We find that in early embryonic development access of maternally deposited nuclear proteins to the genome is temporally ordered via importin affinities.
  36. [36]
    The Maternal to Zygotic Transition in Mammals - PMC - NIH
    The maternal to zygotic transition (MZT) is initiated when sperm and egg fuse at the time of fertilization. Each haploid gamete forms a pronucleus and after ...2.1 Fertilization · 2.2 Zona Pellucida · 3. Epigenetic Regulation Of...
  37. [37]
    De ovi mammalium et hominis genesi (1827), by Karl Ernst von Baer
    Feb 9, 2017 · The 1827 publication provided evidence for the claim that the development of animal life begins from an egg. Sources. Alexandre, Henri. "A ...
  38. [38]
    [PDF] The discovery of meiosis by E. Van Beneden, a breakthrough in the ...
    May 3, 2025 · He investigated the development of the egg in rabbits and bats and summarized his results in a paper published in 1875. The study of.
  39. [39]
    In the beginning… Animal fertilization and sea urchin development
    Dec 1, 2006 · Thus, it was not until Oskar Hertwig published his seminal observations of sperm and egg pronuclear fusion in the sea urchin in 1876 that the ...Missing: timeline | Show results with:timeline
  40. [40]
    An electron microscopic study of sperm penetration into the rabbit ...
    Soon after exposure to ooplasm, the sperm nucleus begins to decondense at a variable rate into a web of electron-dense strands; this process begins in the ...
  41. [41]
    induction of nuclear decondensation of mammalian spermatozoa in ...
    Austin, C.R. (1961) The Mammalian Egg. Thomas, Springfield, Illinois. Bedford, J.M. (1972) An electron microscopic study of sperm penetration into the ...
  42. [42]
    Some reflections on double fertilization, from its discovery to the ...
    Jul 25, 2003 · Double fertilization is a defining feature of reproductive development in the most evolutionarily successful and wonderfully diverse group of ...Missing: pronuclei | Show results with:pronuclei
  43. [43]
    IVF and embryo transfer: historical origin and development
    Some penetration of spermatozoa into the ova, pronuclear formation and cleavage were observed with all media, although at low rates. These experiments did not ...
  44. [44]
    A.R.T. and history, 1678–1978 | Human Reproduction
    ... IVF). The first report on human IVF was by Edwards et al. (1969) who observed pronuclear formation in a few oocytes. In 1970 they reported that they had ...
  45. [45]
    The Role of Ca2+ in Signal Transduction Following Fertilization in ...
    Nov 1, 1993 · W. R.. (. 1993. ). Sources of calcium in sea urchin eggs during the fertilization response . ... fertilization in Pelvetia . Devl Biol . 157.
  46. [46]
    Dynamic alterations in the paternal epigenetic landscape following ...
    Jul 30, 2012 · The majority of protamines (approximately 80%) were removed within 3 h, at which point histone association with DNA begins and is completed by ...
  47. [47]
    CRISPR/Cas9-mediated gene editing in human tripronuclear zygotes
    Apr 18, 2015 · In this report, we used tripronuclear (3PN) zygotes to further investigate CRISPR/Cas9-mediated gene editing in human cells.
  48. [48]
    Atypical centrioles during sexual reproduction - Frontiers
    This review centers on and revolves around the formation, structure, and function of the atypical centrioles found in sexual reproduction.
  49. [49]
    Pronuclear morphology and chromosomal abnormalities as scoring ...
    In humans, the sperm centriole organizes the microtubules, which direct apposition of the two pronuclei and initiate the formation of polar axes at syngamy by ...Missing: timeline | Show results with:timeline<|separator|>
  50. [50]
    Embryo Development - IVF.net
    The pronuclei are usually checked between 16-18 hours after the sperm is added to the egg. 2PN. The 2PN will eventually disappear with the union of two gametes ...
  51. [51]
    Observation of pronuclei may not be an absolute indicator for ... - NIH
    Upon entering the oocyte cytoplasm, the sperm nucleus undergoes a dramatic morphological transformation as the male pronucleus is formed and synthesizes DNA.
  52. [52]
    The Chromosomal Constitution of Embryos Arising from ... - NIH
    Uniparental diploidy may sometimes occur if one pronucleus fails to develop and the other pronucleus already contains a diploid genome or alternatively a ...
  53. [53]
    Time-lapse monitoring of fertilized human oocytes focused on the ...
    Sep 22, 2021 · The appearance of pronuclei (PN) in oocytes is widely used to assess fertilization in human in vitro fertilization (IVF) programs. Oocytes ...
  54. [54]
    Time-lapse imaging: Morphokinetic analysis of in vitro fertilization ...
    Time-lapse imaging (TLI) allows continuous monitoring of embryo development without disturbing culture conditions by removing embryos from the incubator.
  55. [55]
    Three pro-nuclei (3PN) incidence factors and clinical outcomes - NIH
    ... sperm abnormality [24]. In this study, we selected semen of donors to exclude abnormal sperm influence as much as possible in IVF-ET cycles. When 3PN ...
  56. [56]
    Total fertilization failure: is it the end of the story? - PMC - NIH
    ... fertilization (IVF) laboratories, fertilization rates approach 70–80 % [1]. However, fertilization failure still exists as a frustrating experience. Not ...
  57. [57]
    Mutations in PLCZ1 induce male infertility associated with ... - NIH
    Polyspermy is a major cause of multipronuclear fertilization, with up to 80% of multipronuclear fertilization resulting from polyspermic fertilization [5, 6].
  58. [58]
    Preventing polyspermy in mammalian eggs—Contributions of the ...
    Mar 27, 2020 · The egg's blocks to polyspermy (fertilization of an egg by more than one sperm) were originally identified in marine and aquatic species ...
  59. [59]
    Prognostic value of triploid zygotes on intracytoplasmic sperm ...
    Triploidy formation resulting from standard IVF insemination could be due to polyspermy (diandric causes). In the present study, we minimize diandric causes of ...
  60. [60]
    Influencing factors of three pronuclei incidence and their impact on ...
    Multiple mechanisms prevent polyspermy to ensure normal fertilization. However, despite these safeguards, the formation of 3PN zygotes can still occur in some ...
  61. [61]
    Fertilization, Oocyte Activation, Calcium Release and Epigenetic ...
    Mar 4, 2022 · Calcium signaling parameters, particularly amplitude, are important determinants of postfertilization development (Ozil and Huneau, 2001).
  62. [62]
    IVF: Oocyte postmaturity and pronucleus size asynchrony after ICSI
    The cause of fertilization failure after ICSI may be related to ooplasmic immaturity or suboptimal activation capacity of the spermatozoa, both of which result ...Missing: defects | Show results with:defects
  63. [63]
    Defects in phospholipase C zeta cause polyspermy and low ... - NIH
    Mutations in PLCZ1, the gene encoding PLCζ, cause male infertility and intracytoplasmic sperm injection (ICSI) fertilization failure.<|separator|>
  64. [64]
    Assessing the clinical viability of micro 3 pronuclei zygotes - PMC
    The origin of 1PN zygotes may be due to parthenogenetic oocyte activation or asynchronous pronuclear formation and have been linked with implantation failure, ...
  65. [65]
    Novel insights into the genetic and epigenetic paternal contribution ...
    During fertilization, the sperm transmits not only nuclear DNA to the oocyte but also activation factor, centrosomes, and a host of messenger RNA and microRNAs.
  66. [66]
    Pronuclear epigenetic modification of protamine deficient human ...
    Aug 6, 2025 · Epigenetic abnormalities and abnormal chromatin structure in sperm may lead to male infertility. Protamine deficiency is among the disorders ...
  67. [67]
    Assisted reproduction treatment and epigenetic inheritance
    At the level of genomic imprinting involving CpG methylation, ART-induced epigenetic defects are convincingly observed in mice, especially for placenta, and ...
  68. [68]
    The Frequency of Chromosomal Euploidy Among 3PN Embryos - PMC
    In the IVF laboratory, pronuclear status, subsequent development rate and embryo morphology are common parameters in the selection of good quality embryos.Missing: implications | Show results with:implications
  69. [69]
    Association between clinical and IVF laboratory parameters and ...
    Dec 14, 2021 · Several studies have shown that embryo aneuploidy is the main contributing factor to failed IVF, reinforcing the relevance of PGT-A as a means ...Missing: implications | Show results with:implications
  70. [70]
    Polyspermy prevention: facts and artifacts? - PMC - NIH
    Polyspermy is a fact in aged oocytes in vivo and in vitro, whereas the concept of “polyspermy-blocking mechanisms” in mammalian oocytes is mainly based on ...
  71. [71]
    Ion channels and signaling pathways used in the fast polyspermy ...
    Mar 1, 2021 · The fast polyspermy block uses a fertilization-activated depolarization of the egg membrane to electrically inhibit supernumerary sperm from entering the egg.<|control11|><|separator|>