|Embryology - 20 Mar 2019 Expand to Translate|
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- 1 Introduction
- 2 Some Recent Findings
- 3 Spermatozoa Movies
- 4 Seminiferous Tubule
- 5 Spermatozoa Structure
- 6 Human Spermatozoa Development
- 7 Spermatogonia
- 8 Sertoli Cell
- 9 Spermatogenic Cycle
- 10 Spermatozoa Structure
- 11 Meiosis
- 12 Mature Human Spermatozoa
- 13 Spermatozoa Morphology
- 14 Spermatozoa Chemotaxis
- 15 Human Spermatazoa Statistics
- 16 Sertoli Cell
- 17 Histology
- 18 Male Abnormalities
- 19 Additional Images
- 20 References
- 21 Terms
- 22 External Links
- 23 Glossary Links
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)
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.
<pubmed limit=5>Spermatozoa Development</pubmed>
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 - secondary spermatocytes - 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.
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
During infancy and childhood, sertoli cells are the most active cell population in the seminiferous tubule producing AMH from fetal period until mid-puberty.
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 (manchette).
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.
Spermatozoa maturation involves two processes meiosis and spermiogenesis. After puberty, new spermatozoa continue to be generated throughout life from a spermatogonia stem cell population in the testis.
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: Cell Division - Meiosis
Mature Human 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. 20852650
- 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.
Human Spermatazoa Statistics
The following data is based normal human male values for reproductive ages between 20 to 50 years:
- 45 to 207 million spermatozoa produced per day within the two testes
- 2,000 spermatozoa approx per second each day
- Compare this to adult human red blood cell production of about 250,000 million RBCs per day
- 182 million spermatozoa stored (epididymal reserves) up to per epididymis
- 440 million spermatozoa extragonadal stored
- 225 million extragonadal spermatozoa in the ductuli deferentia and caudae epididymides per ejaculation
- 23 million spermatozoa approx (all animals) per gram testicular parenchyma per day
- Transit times
- 0.72 day spermatozoa through the caput
- 0.71 days spermatozoa through the corpus
- 1.76 days spermatozoa through the cauda epididymidis
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.
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
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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- acrosome - Cap-shaped cellular structure formed from the golgi apparatus and contains enzymes to dissolve the oocyte (egg) zona pellucida for fertilisation.
- acrosome compaction - Acrosome reshaping process in final stages of spermatogenesis (spermatid to spermatozoa).
- acrosome reaction - Chemical change within the spermatozoa following binding to the zona pellucida, only acrosome reacted spermatozoa have an ability to fuse with oocytes.
- annulus - Cytoskeletal (septin) structure located between the midpiece and principal piece regions of the tail, thought to form a diffusion barrier between these two domains. PMID 20042538
- asthenozoospermia - (asthenospermia) Term for reduced sperm motility and can be the cause of male infertility.
- axoneme - (axonema) The basic structure in cilia and eukaryotic flagella and in the spermatozoa tail, consisting of parallel microtubules in a characteristic "9 + 2" pattern. This pattern describes 9 outer microtubule doublets (pairs) surrounding 2 central singlet microtubules, in humans 50 μm long. The motor protein dynenin move the outer microtubules with respect to the central pair, bending the cilia and generating motility. Note that prokaryotic bacteria have a similar process (flagellum) that uses an entirely different mechanism for motility.
- blood-testis barrier - (BTB) Formed by tight junctions, basal ectoplasmic specializations, desmosome-like junctions and gap junctions between adjacent Sertoli cells near the basement membrane of the seminiferous epithelium.
- capacitation - term describing the process by which spermaozoa become capable of fertilizing an oocyte, requires membrane changes, removal of surface glycoproteins and increased motility.
- CatSper - cationic (Ca2+) channel of spermatozoa, progesterone activated involved in hyperactivation, acrosome reaction, and possibly chemotaxis.
- centriole - a microtubule organising centre. First required for axoneme formation (distal centriole) that is lost and a second for pronuclei formation (proximal) following fertilisation. Rodents loose both and only have maternal centrioles.
- connecting piece - linkage between the spermatozoa head and the midpiece of the tail. PMID 22767409
- cytoplasmic bridges - Transient cytoplasm connections between spermatids arising from one spermatogonium due to incomplete cytokinesis.
- diploid - (Greek, di = double + ploion = vessel) Having two sets of chromosomes, the normal state for all cells other than the gametes.
- end piece - Last portion of the spermatozoa tail region.
- fibrous sheath - cytoskeletal structure surrounding the axoneme and outer dense fibers, defining the extent of the principal piece region.
- haploid - (Greek, haploos = single) Having a single set of chromosomes as in mature germ/sex cells (oocyte, spermatozoa) following reductive cell division by meiosis. Normally cells are diploid, containing 2 sets of chromosomes.
- interstitial cell - (Leydig cell) Male gonad (testis) cell which secrete the androgen testosterone, beginning in the fetus.
- Johnsen score - a clinical score (1-10) for assessing spermatogenesis in a human testicular biopsy. Named after the author of the original article. PMID 5527187
- Leydig cell - (interstitial cell) Male gonad (testis) cell which secrete the androgen testosterone, beginning in the fetus. These cells are named after Franz von Leydig (1821 - 1908) a German scientist who histologically described these cells.
- meiosis - The cell division that occurs only in production of germ cells where there is a reduction in the number of chromosomes (diploid to haploid) which is the basis of sexual reproduction. All other non-germ cells in the body divide by mitosis.
- midpiece - (middle piece) spermatozoa tail initial segment of axoneme surrounded outer dense fibres then by mitochondria. Next in the tail is the principal piece then finally the end piece.
- mitosis - The normal division of all cells, except germ cells, where chromosome number is maintained (diploid). In germ cell division (oocyte, spermatozoa) meiosis is a modified form of this division resulting in reduction in genetic content (haploid). Mitosis, division of the nucleus, is followed by cytokinesis the division of the cell cytoplasm and the cytoplasmic contents. cytokinesis overlaps with telophase.
- outer dense fibres - (ODF, outer dense fibers) cytoskeletal structures that surround the axoneme in the middle piece and principal piece of the spermatozoa tail.
- primary spermatocyte - arranged in the seminiferous tubule wall deep (luminal) to the spermatogonia. These large cells enter the prophase of the first meiotic division. (More? Meiosis)
- principal piece - Spermatozoa tail segment containing the plasma membrane calcium channels (CatSper1 and CatSper2) required for hyperactivation of motility. Region is partially separated from the midpiece by a barrier called the annulus.
- Sertoli cells - (sustentacular cell) These cells are the spermatozoa supporting cells, nutritional and mechanical, as well as forming a blood-testis barrier. The cell cytoplasm spans all layers of the seminiferous tubule. The cells are named after Enrico Sertoli (1842 - 1910), and italian physiologist and histologist.
- sperm annulus - (Jensen's ring; Latin, annulus = ring) A region of the mammalian sperm flagellum connecting the midpiece and the principal piece. The annulus is a septin-based structure formed from SEPT1, 4, 6, 7 and 12. Septins are polymerizing GTPases that can act as a scaffold forming hetero-oligomeric filaments required for cytokinesis and other cell cycle roles.
- spermatogenesis - (Greek, genesis = origin, creation, generation) The term used to describe the process of diploid spermatagonia division and differentiation to form haploid spermatazoa within the testis (male gonad). The process includes the following cellular changes: meiosis, reoorganization of DNA, reduction in DNA content, reorganization of cellular organelles, morphological changes (cell shape). The final process of change in cell shape is also called spermiogenesis.
- spermatogenesis - (Greek, genesis = origin, creation, generation) The maturation process of the already haploid spermatazoa into the mature sperm shape and organization. This process involves reorganization of cellular organelles (endoplasmic reticulum, golgi apparatus, mitochondria), cytoskeletal changes (microtubule organization) and morphological changes (cell shape, acrosome and tail formation).
- spermatogonia - The cells located in the seminiferous tubule adjacent to the basal membrane that either divide and separate to renew the stem cell population, or they divide and stay together as a pair (Apr spermatogonia) connected by an intercellular cytoplasmic bridge to differentiate and eventually form spermatazoa.
- spermatozoa head - Following spermiogenesis, the first region of the spermatozoa containing the haploid nucleus and acrosome. In humans, it is a flattened structure (5 µm long by 3 µm wide) with the posterior part of nuclear membrane forming the basal plate region. The human spermatozoa is about 60 µm long, actively motile and divided into 3 main regions (head, neck and spermatozoa tail).
- spermatozoa neck - Following spermiogenesis, the second region of the spermatozoa attached to basal plate, transverse oriented centriole, contains nine segmented columns of fibrous material, continue as outer dense fibres in tail. In humans, it forms a short structure (1 µm). The human spermatozoa is about 60 µm long, actively motile and divided into 3 main regions (head, neck and tail).
- spermatozoa tail - Following spermiogenesis, the third region of the spermatozoa that has a head, neck and tail). The tail is also divided into 3 structural regions a middle piece, a principal piece and an end piece. In humans: the middle piece (5 µm long) is formed by axonema and dense fibres surrounded by mitochondria; the principal piece (45 µm long) fibrous sheath interconnected by regularly spaced circumferential hoops; the final end piece (5 µm long) has an axonema surrounded by small amount of cytoplasm and plasma membrane.
- spermatogonial stem cells - (SSCs) The spermatagonia cells located beside the seminiferous tubule basal membrane that either divide and separate to renew the stem cell population, or they divide and stay together as a pair (|Apr spermatogonia) connected by an intercellular cytoplasmic bridge to differentiate and eventually form spermatazoa.
- spermatozoon - singular form of of spermatozoa.
- sperm protein 56 - A component of the spermatozoa acrosomal matrix released to the sperm surface during capacitation.
- teratospermia - Clinical term for a spermatozoa with abnormal morphology (small, large, defects in the head, tail, and/or mid-piece) present in the semen or ejaculate.
See also: Spermatozoa Terms collapse table
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Cite this page: Hill, M.A. (2019, March 20) Embryology Spermatozoa Development. Retrieved from https://embryology.med.unsw.edu.au/embryology/index.php/Spermatozoa_Development
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