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Spermatocytes

Anatomy and Physiology · NLM MeSH · updated 2026-10-31 · version 1

Citations verified against PubMedNot yet clinician-reviewed10 referencesApproximately 3 min read

Male germ cells derived from spermatogonia.

Table of Contents

1. Definition2. Research landscape3. Reading the evidence4. Open-access studies5. Key reviews6. References7. About this article

Definition and research guide

This page combines a MeSH definition, publication statistics and selected research abstracts. It is a starting point for reading the literature, not a medical textbook or a systematic evidence review.

Content available: 4 complete abstracts (571 words), 10 references. Full article text is not stored or reproduced here; follow the PMID, DOI or PMC links to read the original sources and their access terms.

Definition

Male germ cells derived from spermatogonia. The euploid primary spermatocytes undergo meiosis and give rise to the haploid secondary spermatocytes which in turn give rise to spermatids.

Definition: NLM Medical Subject Headings (MeSH) scope note (Public domain (U.S. National Library of Medicine); attribution requested). View the MeSH record.

Also known as: Spermatocyte; Spermiocytes; Spermiocyte.

Where it fits in the MeSH classification

Broader topics
Spermatozoa
See also
Sperm Maturation

Related concepts

  • Spermatogonia
  • Meiosis
  • Spermatids

Research landscape

PubMed indexes 4,411 records tagged with the MeSH heading “Spermatocytes” (counts retrieved 2026-10-31). 477 of them (10.8%) were published from 2020 onward, and the busiest period so far is through 1999 with 1,839 records.

Reviews account for 144 records (3.3%) and meta-analyses for 0. Randomized controlled trials number 0 (0.0%) as a share of all records on the topic. Randomized trials make up only a small share of this literature, so much of what is known comes from laboratory, observational or descriptive work.

PubMed records by publication periodthrough 1999: 1,839 records; 2000-2009: 1,038 records; 2010-2019: 1,057 records; 2020 onward: 477 records1,839through 19991,0382000-20091,0572010-20194772020 onward
Records in PubMed indexed with this heading, by publication period. Real counts from NCBI E-utilities; the final period covers only part of the current year. Volume reflects how much has been published, not how strong the evidence is.
Show the data
PeriodRecords
through 19991,839
2000-20091,038
2010-20191,057
2020 onward477
Share of records by publication typeReviews: 3.3 percent; Meta-analyses: 0.0 percent; Randomized controlled trials: 0.0 percentReviews3.3% (144)Meta-analyses0.0% (0)Randomized controlled trials0.0% (0)
Share of all PubMed records on this heading that carry each publication type. Types overlap with other records, so the shares do not sum to 100%.
Publication-type chart data
Publication types among 4,411 PubMed records
TypeRecordsShare
Reviews1443.3%
Meta-analyses00.0%
Randomized controlled trials00.0%

Method: PubMed query "Spermatocytes"[MeSH Terms] with publication-type and date filters, retrieved 2026-10-31.

Reading the evidence in anatomy and physiology

Anatomy describes how body structures are built and physiology describes how they normally work, so most findings here are descriptive or mechanistic rather than evidence about treatment. Much of what is known comes from dissection, imaging, tissue studies and experiments in animals, and measurements in healthy volunteers can differ from those in people with disease or at different ages. Normal ranges vary between individuals and populations. Check whether a result was measured in people or in another species, the age and health of the participants, and the method used.

Evidence from open-access studies

These open-access studies come from PubMed’s best-match results for this heading (papers whose main topic it is), ranked here by citation count, with one recent addition. Each abstract is reproduced in full under its recorded open licence so that you can read the authors’ own words. They illustrate the literature; they are not a systematic summary of it. Citation counts measure attention, not reliability or clinical benefit.

Study comparison — selected abstracts, not a systematic review
Study / authorsYearSource label / designEurope PMC citationsLicence / source
Staged developmental mapping and X chromosome transcriptional dynamics during mouse spermatogenesis
Ernst C, Eling N, Martinez-Jimenez CP, Marioni JC, Odom DT
2019Research article
Design not recorded in structured metadata
247cc by
PMID 30890697 · DOI · PMC6424977
Testis single-cell RNA-seq reveals the dynamics of de novo gene transcription and germline mutational bias in Drosophila
Witt E, Benjamin S, Svetec N, Zhao L
2019Research article
Design not recorded in structured metadata
157cc by
PMID 31418408 · DOI · PMC6697446
ATM and PRDM9 regulate SPO11-bound recombination intermediates during meiosis
Paiano J, Wu W, Yamada S, Sciascia N, Callen E, Paola Cotrim A, et al.
2020Research article
Design not recorded in structured metadata
107cc by
PMID 32051414 · DOI · PMC7016097
Dysregulation of N-glycosylation by Rpn1 knockout in spermatocytes induces male infertility via endoplasmic reticulum stress in mice
Zhang M, Li M, Li H, Wan Y, Yang S, Ji S, et al.
2025Research article
Design not recorded in structured metadata
7cc by
PMID 40083683 · DOI · PMC11900820

Staged developmental mapping and X chromosome transcriptional dynamics during mouse spermatogenesis [1]

Research article Ernst C, Eling N, Martinez-Jimenez CP, Marioni JC, Odom DT · · 2019 · cited 247 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
Male gametes are generated through a specialised differentiation pathway involving a series of developmental transitions that are poorly characterised at the molecular level.
Closing excerpt
This identifies a set of genes strongly repressed by H3K9me3 in spermatocytes, which then undergo extensive chromatin remodelling post-meiosis, thus acquiring an active chromatin state and spermatid-specific expression.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (142 words)

Male gametes are generated through a specialised differentiation pathway involving a series of developmental transitions that are poorly characterised at the molecular level. Here, we use droplet-based single-cell RNA-Sequencing to profile spermatogenesis in adult animals and at multiple stages during juvenile development. By exploiting the first wave of spermatogenesis, we both precisely stage germ cell development and enrich for rare somatic cell-types and spermatogonia. To capture the full complexity of spermatogenesis including cells that have low transcriptional activity, we apply a statistical tool that identifies previously uncharacterised populations of leptotene and zygotene spermatocytes. Focusing on post-meiotic events, we characterise the temporal dynamics of X chromosome re-activation and profile the associated chromatin state using CUT&RUN. This identifies a set of genes strongly repressed by H3K9me3 in spermatocytes, which then undergo extensive chromatin remodelling post-meiosis, thus acquiring an active chromatin state and spermatid-specific expression.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 30890697 · DOI · PMC6424977

Testis single-cell RNA-seq reveals the dynamics of de novo gene transcription and germline mutational bias in Drosophila [2]

Research article Witt E, Benjamin S, Svetec N, Zhao L · · 2019 · cited 157 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
The testis is a peculiar tissue in many respects.
Closing excerpt
By elucidating the distribution of genetic novelties across spermatogenesis, this study provides a deeper understanding of how the testis maintains its core reproductive function while being a hotbed of evolutionary innovation.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (150 words)

The testis is a peculiar tissue in many respects. It shows patterns of rapid gene evolution and provides a hotspot for the origination of genetic novelties such as de novo genes, duplications and mutations. To investigate the expression patterns of genetic novelties across cell types, we performed single-cell RNA-sequencing of adult Drosophila testis. We found that new genes were expressed in various cell types, the patterns of which may be influenced by their mode of origination. In particular, lineage-specific de novo genes are commonly expressed in early spermatocytes, while young duplicated genes are often bimodally expressed. Analysis of germline substitutions suggests that spermatogenesis is a highly reparative process, with the mutational load of germ cells decreasing as spermatogenesis progresses. By elucidating the distribution of genetic novelties across spermatogenesis, this study provides a deeper understanding of how the testis maintains its core reproductive function while being a hotbed of evolutionary innovation.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 31418408 · DOI · PMC6697446

ATM and PRDM9 regulate SPO11-bound recombination intermediates during meiosis [3]

Research article Paiano J, Wu W, Yamada S, Sciascia N, Callen E, Paola Cotrim A, et al. · · 2020 · cited 107 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
Meiotic recombination is initiated by SPO11-induced double-strand breaks (DSBs).
Closing excerpt
Thus, in addition to governing SPO11 breakage, ATM and PRDM9 are critical local regulators of mammalian SPO11 processing.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (148 words)

Meiotic recombination is initiated by SPO11-induced double-strand breaks (DSBs). In most mammals, the methyltransferase PRDM9 guides SPO11 targeting, and the ATM kinase controls meiotic DSB numbers. Following MRE11 nuclease removal of SPO11, the DSB is resected and loaded with DMC1 filaments for homolog invasion. Here, we demonstrate the direct detection of meiotic DSBs and resection using END-seq on mouse spermatocytes with low sample input. We find that DMC1 limits both minimum and maximum resection lengths, whereas 53BP1, BRCA1 and EXO1 play surprisingly minimal roles. Through enzymatic modifications to END-seq, we identify a SPO11-bound meiotic recombination intermediate (SPO11-RI) present at all hotspots. We propose that SPO11-RI forms because chromatin-bound PRDM9 asymmetrically blocks MRE11 from releasing SPO11. In Atm-/- spermatocytes, trapped SPO11 cleavage complexes accumulate due to defective MRE11 initiation of resection. Thus, in addition to governing SPO11 breakage, ATM and PRDM9 are critical local regulators of mammalian SPO11 processing.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 32051414 · DOI · PMC7016097

Dysregulation of N-glycosylation by Rpn1 knockout in spermatocytes induces male infertility via endoplasmic reticulum stress in mice [4]

Research article Zhang M, Li M, Li H, Wan Y, Yang S, Ji S, et al. · · 2025 · cited 7 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
N-glycosylation protein modification plays a crucial regulatory role in numerous biological processes, although their contribution to male reproduction in mammals remains largely undefined.
Closing excerpt
These findings highlight the essential physiological functions of N-glycosylation modification in male spermatogenesis and expand our understanding of its role in male fertility.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (131 words)

N-glycosylation protein modification plays a crucial regulatory role in numerous biological processes, although their contribution to male reproduction in mammals remains largely undefined. Here, we found that Ribophorin I (RPN1), a subunit of oligosaccharyltransferase complex, is indispensable for spermatogenesis in male germ cells. Germ cell-specific Rpn1 knockout results in significant inhibition of the progression of meiosis, consequently disrupting homologous chromosome pairing, meiotic recombination, and DNA double strand breaks repair during meiosis. N-glycoproteomic profiling revealed that glycosylation levels are reduced in endoplasmic reticulum-associated proteins, while functional analyses showed that Rpn1 deficiency could inhibit endoplasmic reticulum function and trigger endoplasmic reticulum stress during meiosis and increasing apoptosis levels in mice. These findings highlight the essential physiological functions of N-glycosylation modification in male spermatogenesis and expand our understanding of its role in male fertility.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 40083683 · DOI · PMC11900820

Key reviews and meta-analyses

The most cited review-type papers among PubMed’s best-match results for this heading, listed with bibliographic details only. Follow the links for the full text and access terms.

  • Research article Sex matters in meiosis [5]
    Hunt PA, Hassold TJ · Science · 2002 · cited 291 times (Europe PMC)
  • Research article Regulation of spermatogonial stem cell self-renewal and spermatocyte meiosis by Sertoli cell signaling [6]
    Chen SR, Liu YX · Reproduction · 2015 · cited 226 times (Europe PMC)
  • Research article A possible meiotic function of the peculiar patterns of gene expression in mammalian spermatogenic cells [7]
    Kleene KC · Mech Dev · 2001 · cited 174 times (Europe PMC)
  • Research article Surfing the wave, cycle, life history, and genes/proteins expressed by testicular germ cells. Part 1: background to spermatogenesis, spermatogonia, and spermatocytes [8]
    Hermo L, Pelletier RM, Cyr DG, Smith CE · Microsc Res Tech · 2010 · cited 145 times (Europe PMC)
  • Research article Surfing the wave, cycle, life history, and genes/proteins expressed by testicular germ cells. Part 2: changes in spermatid organelles associated with development of spermatozoa [9]
    Hermo L, Pelletier RM, Cyr DG, Smith CE · Microsc Res Tech · 2010 · cited 139 times (Europe PMC)
  • Research article Genetics of germ cell development [10]
    Lesch BJ, Page DC · Nat Rev Genet · 2012 · cited 115 times (Europe PMC)

References

  1. Ernst C, Eling N, Martinez-Jimenez CP, Marioni JC, Odom DT. Staged developmental mapping and X chromosome transcriptional dynamics during mouse spermatogenesis. . 2019. PMID 30890697 · DOI · PMC6424977
  2. Witt E, Benjamin S, Svetec N, Zhao L. Testis single-cell RNA-seq reveals the dynamics of de novo gene transcription and germline mutational bias in Drosophila. . 2019. PMID 31418408 · DOI · PMC6697446
  3. Paiano J, Wu W, Yamada S, Sciascia N, Callen E, Paola Cotrim A, et al.. ATM and PRDM9 regulate SPO11-bound recombination intermediates during meiosis. . 2020. PMID 32051414 · DOI · PMC7016097
  4. Zhang M, Li M, Li H, Wan Y, Yang S, Ji S, et al.. Dysregulation of N-glycosylation by Rpn1 knockout in spermatocytes induces male infertility via endoplasmic reticulum stress in mice. . 2025. PMID 40083683 · DOI · PMC11900820
  5. Hunt PA, Hassold TJ. Sex matters in meiosis. Science. 2002. PMID 12077403 · DOI
  6. Chen SR, Liu YX. Regulation of spermatogonial stem cell self-renewal and spermatocyte meiosis by Sertoli cell signaling. Reproduction. 2015. PMID 25504872 · DOI
  7. Kleene KC. A possible meiotic function of the peculiar patterns of gene expression in mammalian spermatogenic cells. Mech Dev. 2001. PMID 11472831 · DOI
  8. Hermo L, Pelletier RM, Cyr DG, Smith CE. Surfing the wave, cycle, life history, and genes/proteins expressed by testicular germ cells. Part 1: background to spermatogenesis, spermatogonia, and spermatocytes. Microsc Res Tech. 2010. PMID 19941293 · DOI
  9. Hermo L, Pelletier RM, Cyr DG, Smith CE. Surfing the wave, cycle, life history, and genes/proteins expressed by testicular germ cells. Part 2: changes in spermatid organelles associated with development of spermatozoa. Microsc Res Tech. 2010. PMID 19941292 · DOI
  10. Lesch BJ, Page DC. Genetics of germ cell development. Nat Rev Genet. 2012. PMID 23044825 · DOI

About this article

Sources
NLM Medical Subject Headings (definition, synonyms); NCBI PubMed (counts, identifiers); Europe PMC (open-access abstracts and citation counts).
Verification
Every reference was resolved against PubMed on 2026-10-31; title, year and DOI matched.
Review state
automated source verified; no human expert review has been recorded yet.
Version
1, last updated 2026-10-31.

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Related research topics

  • SpermatidsAnatomy and Physiology
  • SpermatogoniaAnatomy and Physiology
  • SpermatozoaAnatomy and Physiology
  • AcrosomeAnatomy and Physiology
  • Sperm HeadAnatomy and Physiology
  • OocytesAnatomy and Physiology

All 542 articles in Anatomy and Physiology →

At a glance

4,411
PubMed records on this heading
11%
published since 2020
144
review articles
0
randomized trials
10
verified references below

Counts come from PubMed on 2026-10-31; they show how much has been written, not how strong the evidence is.

Related topics See all →

  • SpermatidsAnatomy and Physiology
  • SpermatogoniaAnatomy and Physiology
  • SpermatozoaAnatomy and Physiology
  • AcrosomeAnatomy and Physiology
  • Sperm HeadAnatomy and Physiology

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Sources

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  • PubMed search used ↗

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