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This page introduces spermatogenesis the development of spermatozoa, the male haploid gamete cell. In humans at puberty, spermatozoa are produced by spermatogonia meiosis in the seminiferous tubules of the testis (male gonad). A second process of spermiogenesis leads to change in cellular organisation and shape before release into the central lumen of the seminiferous tubule. This overall process has been variously divided into specific identifiable stages in different species: 6 in human, 12 in mouse, and 14 in rat. Structurally, the seminiferous tubule epithelium is divided into a basal and an apical (adluminal) compartment by the blood–testis barrier (BTB). (More? Testis Development).
A second unique feature of this process is that during mitosis and meiosis the dividing cells remain connected by cytoplasmic bridges as the cells do not complete cytokinesis. This cellular organization is described as a syncytium, only ending with release into the central lumen of the seminiferous tubule, when the cell cytoplasm is discarded. Retinoic acid has been shown to be a key regulator of the development process. (More? Retinoic acid)
- In a healthy adult human male it takes about 48 days from meiosis to produce a mature spermatozoa, and he produces somewhere between 45 to 207 million spermatozoa per day, or about 1 to 2,000 every second. (More? Statistics)
|Genital Links: genital | Lecture - Medicine | Lecture - Science | Lecture Movie | Medicine - Practical | primordial germ cell | meiosis | endocrine gonad | Genital Movies | genital abnormalities | Assisted Reproductive Technology | puberty | Category:Genital|
Some Recent Findings
|More recent papers|
This table allows an automated computer search of the external PubMed database using the listed "Search term" text link.
Search term: Spermatozoa Development | Spermatogonia Development | Spermatogonia Stem Cells | Spermatozoa Meiosis | Primary Spermatocyte | Secondary Spermatocyte | Spermatid | Spermatogenesis | Spermiogenesis | Sertoli Cell Development | Spermatozoa Chemotaxis | Acrosome
|These papers originally appeared in the Some Recent Findings table, but as that list grew in length have now been shuffled down to this collapsible table.
See also Week 1 movies.
Seminiferous tubule cartoon
- Spermatogonia - are the first cells of spermatogenesis
- Primary spermatocyte - large, enter the prophase of the first meiotic division
- Secondary spermatocytes - small, complete the second meiotic division
- Spermatid - immature spermatozoa
- Spermatozoa - differentiated gamete
- Spermatozoa development: primordial germ cell - spermatogonia - primary spermatocyte - Template:Secondary spermatocyte - spermatid - spermatozoa
Spermatozoa (mouse) cross-sections of tail (EM) and diagram
Other main cell types seen in the histological sections
- sertoli cells- support cells seen within the seminiferous tubule
- Interstitial cells or Leydig cells - produce hormone
- Smooth muscle - surround seminiferous tubule and contribute to contraction of the tubule
Human Spermatozoa Development
- Spermatogenesis process of spermatagonia mature into spermatazoa (sperm).
- Continuously throughout life occurs in the seminiferous tubules in the male gonad- testis (plural testes).
- At puberty spermatagonia activate and proliferate (mitosis).
- about 48 days from entering meiosis until morphologically mature spermatozoa
- about 64 days to complete spermatogenesis, depending reproduction time of spermatogonia
- follicle stimulating hormone (FSH) - stimulates the spermatogenic epithelium
- luteinizing-hormone (LH) - stimulates testosterone production by Leydig cells
In humans at about 2 months of age, primordial germ cells (gonocytes) are replaced by adult dark (Ad) and pale (Ap) spermatogonia forming the spermatogonial stem cell (SSC) population that at puberty will commence differentiation into spermatozoa.
The spermatogonia are the diploid stem cell (spermatogonial stem cell, SSC) progenitor for spermatozoa development. They are located on the basal lamina around the periphery of the seminiferous tubule wall. See this recent spermatogonia review.
In 1963 Clermont identified spermatogonia as Ap (pale) and Ad (dark) on basis of light microscope staining.
- now also type B
- 60 years - Ap spermatogonia number decrease
- 80 years - Ad spermatogonia number decrease
|Containing only spermatogonia and sertoli cells||Containing spermatogonia, sertoli cells and stages of spermatozoa cell meiosis|
|Mouse spermatogonia have been shown to require a number of factors to regulate both their spermatogonial self-renewal and differentiation.Zhou Q & Griswold MD. (2008). Regulation of spermatogonia. , , . PMID: 20614596 DOI.
The mouse "As model" originally stated that the As spermatogonia are the SSCs. A more recent proposal suggests that only some of the As spermatogonia have the potential for long-term self-renewal, while others have a limited capacity, indicating the presence of a SSC hierarchy.
In the mouse testis, spermatogonial stem cells can also be identified by Id4 expression, a dominant-negative transcription factor containing a basic helix-loop-helix (bHLH) region. Id4 inhibits binding to DNA and transcriptional transactivation by hetero-dimerization with other bHLH proteins.
Spermatozoa maturation involves two processes meiosis and spermiogenesis. After puberty, new spermatozoa continue to be generated throughout life from a spermatogonia stem cell (SSC) population in the testis.
The first large differentiating diploid (2N) cell before meiosis I, that enters the prophase of the first meiotic division.
- Search PubMed: Primary Spermatocyte
The primary spermatocyte forms two second small cells that complete the second meiotic division to be haploid (N) . Histologically difficult to observe, these cells will complete meiosis, forming next the spermatid stage that is the immature morphological state of the spermatozoa.
- Search PubMed: Secondary Spermatocyte
Differences in Mammalian Meioses
|Female Oogenesis||Male Spermatogenesis|
|Meiosis initiated||once in a finite population of cells||continuously in mitotically dividing stem cell population|
|Gametes produced||1 / meiosis||4 / meiosis|
|Meiosis completed||delayed for months or years||completed in days or weeks|
|Meiosis Arrest||arrest at 1st meiotic prophase||no arrest differentiation proceed continuously|
|Chromosome Equivalence||All chromosomes exhibit equivalent transcription and recombination during meiotic prophase||Sex chromosomes excluded from recombination and transcription during first meiotic prophase|
|Gamete Differentiation||occurs while diploid (in first meiotic prophase)||occurs while haploid (after meiosis ends)|
- Links: meiosis
Spermiogenesis is the final stage of spermatogenesis, morphological changes transform the round spermatids into the mature spermatozoa shape and structure.
- Nuclear compression - chromatin condensation occurs by the replacement of histones with protamines.
- Acrosome formation - located over the anterior part of the spermatid nucleus, cap-like membrane-bound organelle formed through coalescence of the coated vesicles budding from the trans-Golgi network. The acrosome-acroplaxome-manchette complex is a major driver for the shaping of the spermatozoa head.
- Tail development - located over the posterior part of the spermatid nucleus, initially a centriole pair moves, the axoneme develops from the distal centriole. Axoneme consists of a central pair of microtubules surrounded by 9 outer doublet microtubules ("9 × 2 + 2").
- Cytoplasm disposal - cytoplasm transported towards the tail along the manchette and finally its removal.
- Nuclear pore redistribution - with packaging of the nuclear pores into the redundant and discarded nuclear envelope. 
- Autophagy - a self-digestion process, may also occur regulating cytoskeleton reorganization.
The sertoli cell was named after Enrico Sertoli (1842 - 1910) an Italian (Milan) physiologist and histologist. These cells support spermatozoa development and span the wall of the seminiferous tubule.
- sustentacular cells of seminiferous tubules.
- form a “blood-testis” barrier through junctional complexes
- separate the intra-tubular germinal epithelium into two compartments
- basal compartment - cells are exposed to the extra-tubular environment
- luminal compartment - cells are subject to an environment produced by Sertoli cells and germ cells
Along the length of the seminiferous tubule spermatozoa develop in a cyclic manner over time progressing through a number of stages, called the spermatogenic cycle, see review. The number of stages appears to differ between species, in mouse there are 12 stages (I – XII) and in the rat 14 stages.
In mouse, one spermatogenic cycle (12 stages) occurs over 8.6 days and four cycles (35 days) are required from spermatogonial stem cell to released spermatozoa.
Spermatozoa (mouse) cross-sections of tail (EM) and diagram
This structure forms the acrosome plate with intermediate filament bundles of the marginal ring at the leading edge of the acrosome. The acroplaxome site for Golgi-derived proacrosomal vesicles to tether and fuse and anchors the developing acrosome to the elongating spermatid head and may provide a scaffolding for the shaping of the spermatid nucleus.
Derived from the Golgi apparatus in conjunction with transient specialized bundles of microtubules (Template:Manchette), this vesicle releases its contents following progesterone stimulus or zona pellucida binding.
The "acrosome reaction" (AR) a type of specialised exocytosis, or similar to the release of an exosome. This also leads to changes in the spermatozoa membrane. The 26S proteasome, has been identified from in vitro studies to be required for zona lysin in many species, including mammals.
- Calcium ion permeability - Ca(2+) release from the acrosome leads to exocytosis of the acrosomal vesicle, alkalization appears to be a critical step.
- store-operated Ca(2+) channels and voltage-dependent Ca(2+) channels
- Vesicle membrane - initially holding excreting molecules, remains on the cell surface
- Membrane loss - both outer acrosomal membrane and plasma membrane are lost by forming vesicles during acrosome reaction.
Acrosome reaction has a slow and rapid component:
- Rapid - (seconds) efflux of calcium from intracellular stores, triggers fusion pores opening and the release of hybrid vesicles.
- Slow - (minutes) acrosomal swelling, triggered by activation of an adenylyl cyclase downstream of the opening of store-operated calcium channels. Determines the kinetics of the acrosome reaction.
- Exosomes - extracellular vesicles that are released from cells upon fusion of an intermediate endocytic compartment, the multivesicular body (MVB), with the plasma membrane.
- Proteasomes - protein complexes which typically degrade unneeded or damaged proteins by proteolysis.
- Acrosin is essential for sperm penetration through the zona pellucida in hamsters PNAS February 4, 2020 117 (5) 2513-2518 "Mammalian oocytes are surrounded by the zona pellucida, a glycoprotein coat that protects the oocyte and embryo from mechanical damage during their preimplantation development within the oviduct. Fertilizing spermatozoa must penetrate the zona, but we do not know the exact mechanisms underlying this process. Sperm proteases were thought to work as zona lysins, but gene-knockout studies in mice did not support this assumption. In this study, we generated hamsters without acrosin, the major acrosomal protease, to examine its role in both in vivo and in vitro fertilization. Surprisingly, mutant male hamsters were completely infertile because their spermatozoa were unable to penetrate the zona. We thus demonstrated that, at least in hamsters, acrosin is essential for sperm penetration through the zona."
Human acrosin - 22q13.33
- Links: OMIM - Acrosin
The spermatozoa nucleus undergoes extensive compression, and nuclear DNA chromatin remodelling by tightly packing with spermatozoa-specific protamines.
|It is thought that the lysine-rich protein precursor (H1 histone) has evolved into the arginine-rich protamines.||Three major spermatozoa nuclear basic proteins types:
|EM Human Spermatozoa Nucleus|
|Cap-phase spermatid nucleus||Elongated spermatid nucleus|
|Normal human spermatozoa||Abnormal human spermatozoa|
The stable mature microtubule-containing tail of the sperm.
|Historic EM spermatozoon tail||Mouse cross-sections of tail|
Spermatozoa initially contains 2 centrioles (proximal, distal) and at fertilisation only a single (proximal) is present, which in most mammalian species is contributed to reconstitute the zygotic centrosome. Note that in rodents (rat, mice) both centrioles are lost and only a maternal centrosomal inheritance occurs.
- distal centriole - (perpendicular to membrane) required as the basal body generating the microtubule axoneme and is then lost (disintegration).
- proximal centriole - required after fertilisation for decondensing spermatozoa nucleus allowing development into the male pronucleus.
A transient microtubule structure formed in spermatids involved in the process of: assembly of the mammalian sperm tail, mechanical shaping and condensation of the sperm nucleus. These microtubules are also invloved with specific transport, intramanchette transport, which has been likened to intraflagellar transport. This microtubular structure surrounds the nucleus of the developing spermatid and is thought also to assist in both the reshaping of the nucleus and redistribution of spermatid cytoplasm.
Contained in the initial segment provide the energy for motility and may also enter the egg on fertilization, but are eliminated by a ubiquitin-dependent mechanism.
Located in the sperm head perinuclear region and contains a cytoskeletal element to maintain the shape of the sperm head and functional molecules leading to oocyte activation during fertilization.
Mature Human Spermatozoa
Human Spermatozoa Statistics
|produced / day (two testes)||45 to 207 million|
|compare adult human red blood cell / day||250,000 million|
|produced / second each day (approx)||2,000|
|stored (epididymal reserves) up to per epididymis||182 million|
|stored extragonadal||440 million|
|extragonadal - ductuli deferentia and caudae epididymides per ejaculation||225 million|
|through the caput||0.72 day|
|through the corpus||0.71 days|
|through the cauda epididymidis||1.76 days|
|Table Data  See also WHO human semen reference values(2010). Links: spermatozoa|
Morphology is a term used to describe the overall appearance of a cell or tissue and is often used to characterise changes in cellular state or activity. Historically, there have been studies comparing the overall appearance of spermatozoa between different species. More recently, there have been several different ways of characterising the morphology of human spermatozoa developed mainly in relation to clinical reproductive technologies.
Integrated Sperm Analysis System (ISAS)
A semi-automated computer-aided system that measures spermatozoa head parameters length (L), width (W), area (A), perimeter (P), acrosomal area (Ac), and the derived values L/W and P/A.
- For each man a homogeneous population of distributions characterized seminal spermatozoa (7,942 cells: median values L 4.4 μm, W 2.8 μm, A 9.8 μm(2), P 12.5 μm, Ac 47.5%, L/W 1.57, P/A 1.27)
- Different men could have spermatozoa of significantly different dimensions.
- Head dimensions for swim-up spermatozoa from different men (4 812 cells) were similar to those in semen, differing only by 2%-5%.
- The values of L, W and L/W fell within the limits given by the World Health Organization (WHO).
- A subpopulation of 404 spermatozoa considered to fit the stringent criteria of WHO 'normal' seminal spermatozoa from both semen and swim-up were characterized by median values (and 95% confidence intervals) of L, 4.3 μm (3.8-4.9), W, 2.9 μm (2.6-3.3), A, 10.2 μm(2) (8.5-12.2), P, 12.4 μm (11.3-13.9), Ac, 49% (36-60), L/W, 1.49 (1.32-1.67) and P/A, 1.22 (1.11-1.35). These median values fall within the 95th centile confidence limits given by WHO, but the confidence intervals for L and W were larger.
Chemotaxis was first identified in marine species, which still remains today as a model system. While the signals may differ, the overall effect is to chemically attack spermatozoa to the oocyte to allow fertilisation to occur.
|The following series of 2011 research articles have identified the spermatozoa calcium channel protein (CatSper) as the progesterone activated pathway involved in capacitated spermatozoa chemotaxis.
Human Spermatozoa Chemotaxis Model (2009)
See also 2008 review.
The sertoli cells are the first cells to be differentiated in development by SRY expression. Post-puberty these are the "support" cells for spermatozoa development and transport from the periphery to lumen of the seminiferous tubule. Sertoli cells form a barrier with cell junctions at the Sertoli cell-cell and Sertoli-germ cell interface.
Sertoli cell postnatal proliferation may be regulated by thyroid status. An animal model study of postnatal transient hypothyroidism has demonstrated Sertoli cell proliferation (6 to 8 fold increase) 2 days after the diet switch and remained elevated the next days.
Papanicolaou stain (Papanicolaou's stain, Pap stain) a multichromatic (five dyes) staining histological technique developed by George Papanikolaou, used to differentiate cells in smear preparations of various bodily secretions.
- Links: sertoli cell | Testis Histology | Histology Stains | Search PubMed - Sertoli Cell Development
Male Infertility Genes
Examples of known genes resulting in various forms of male infertility.
|Gene abbreviation||Name||Gene Location||Online Mendelian
Inheritance in Man (OMIM)
|HUGO Gene Nomenclature
|AURKC||Aurora kinase C||19q13.43||603495||11391||GC19P057230||Macrozoospermia|
|CATSPER1||Cation channel sperm-associated 1||11q13.1||606389||17116||GC11M066034||Asthenozoospermia|
|CFTR||Cystic fibrosis transmembrane conductance regulator||7q31.2||602421||1884||GC07P117465||Obstructive azoospermia|
|DNAH1||Dynein axonemal heavy chain 1||3p21.1||603332||2940||GC03P052350||Asthenozoospermia|
|DPY19L2||Dpy-19-like 2 gene||12q14.2||613893||19414||GC12M063558||Globozoospermia|
|GALNTL5||Polypeptide N-acetylgalactosaminyltransferase-like 5||7q36.1||615133||21725||GC07P151956||Asthenozoospermia|
|MAGEB4||MAGE family member B4||Xp21.2||300153||6811||GC0XP030260||Azoospermia|
|NANOS1||Nanos C2HC-type zinc finger 1||10q26.11||608226||23044||GC10P119029||Azoospermia|
|NR0B1||Nuclear receptor subfamily 0 group B member 1||Xp21.2||300473||7960||GC0XM030322||Azoospermia|
|NR5A1||Nuclear receptor subfamily 5 group A member 1||9q33.3||184757||7983||GC09M124481||Azoospermia|
|SOHLH1||Spermatogenesis and oogenesis-specific basic helix–loop–helix 1||9q34.3||610224||27845||C09M135693||Azoospermia|
|SYCE1||Synaptonemal complex central element protein 1||10q26.3||611486||28852||GC10M133553||Azoospermia|
|TAF4B||TATA-box binding protein-associated factor 4b||18q11.2||601689||11538||GC18P026225||Azoospermia|
|TEX11||Testis expressed 11||Xq13.1||300311||11733||GC0XM070528||Azoospermia|
|TEX15||Testis expressed 15, meiosis and synapsis associated||8p12||605795||11738||GC08M030808||Azoospermia|
|WT1||Wilms tumour 1||8p12||607102||12796||GC11M032365||Azoospermia|
|ZMYND15||Zinc-finger MYND-type containing 15||17p13.2||614312||20997||GC17P004740||Azoospermia|
| Table data source (table 1) Links: fertilization | spermatozoa | testis | Male Infertility Genes | Female Infertility Genes | oocyte | ovary | Genetic Abnormalities | ART|
Asthenozoospermia - (asthenospermia) term for reduced spermatozoa motility. Azoospermia - term for no spermatozoa located in the ejaculate. Globozoospermia - term for spermatozoa with a round head and no acrosome.
Human sperm pathologies These electron micrographs show a range of tail structural abnormalities including (a) two tails. (b) shows a normal spermatozoa tail cross-section and c to g show a range of abnormal tail structures (open image to see details).
A clinical score (1-10) for assessing spermatogenesis in a human testicular biopsy. Named after the author of the original article .
|10||complete spermatogenesis and perfect tubules|
|9||many spermatozoa present but disorganized spermatogenesis|
|8||only a few spermatozoa present|
|7||no spermatozoa but many spermatids present|
|6||only a few spermatids present|
|5||no spermatozoa or spermatids present but many spermatocytes present|
|4||only a few spermatocytes present|
|3||only spermatogonia present|
|2||no germ cells present|
|1||neither germ cells nor Sertoli cells present|
Reference: Johnsen SG. Testicular biopsy score count - a method for registration of spermatogenesis in human testes: normal values and results in 335 hypogonadal males. (1970) Hormones 1(1): 2-25. PubMed 5527187
|Severe oligozoospermia||less than 1|
|Normal||greater than 20|
(Low Sperm Count) less than 20 million sperm after 72 hour abstinence from sex
(Absent Sperm) blockage of duct network
Immotile Cilia Syndrome
Lack of sperm motility
Acephalic spermatozoa syndrome
Acephalic spermatozoa syndrome is characterized by the presence of very few intact spermatozoa and tailless sperm heads in the semen and leads to severe male infertility. Sad1 and UNC84 domain-containing 5 (SUN5) is a testis-specific nuclear envelope protein. A recent study has shown that mutations in SUN5 appear to affect the secondary structure of the protein and influence its folding and cellular localization.
- Links: OMIM - SUN5
Infertility - Stem Cells
Recent studies have been able to transplant of own cryostored spermatogonial stem cells (SSCs) is a promising technique for fertility restoration when the SSC pool has been depleted.
Human spermatozoa acrosomal protein SP-10
Human spermatid electron micrograph
Model capacitation-induced acrosome docking to sperm membrane
Mouse spermiogenesis model
Mouse- seminiferous tubule histology
Rat Spermatogenesis figure
Human spermatozoa - phospholipase C zeta localization
Labeled Chemotaxis Model
Spermatogenesis androgen action
Mouse spermatogenesis stages
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|Spermatozoa Development (expand to see terms)|
See also: Spermatozoa Terms collapse table
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- World Health Organization - WHO Laboratory Manual for the Examination and Processing of Human Semen. 5th ed. Geneva, Switzerland: World Health Organization; 2010. Online PDF
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