Male Gamete
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Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, which are known as spermatozoa, while some
red algae Red algae, or Rhodophyta (, ; ), are one of the oldest groups of eukaryotic algae. The Rhodophyta also comprises one of the largest phyla of algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing. The majority ...
and fungi produce non-motile sperm cells, known as spermatia. Flowering plants contain non-motile sperm inside
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
, while some more basal plants like ferns and some gymnosperms have motile sperm. Sperm cells form during the process known as
spermatogenesis Spermatogenesis is the process by which haploid spermatozoa develop from germ cells in the seminiferous tubules of the testis. This process starts with the mitotic division of the stem cells located close to the basement membrane of the tubule ...
, which in
amniote Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are disti ...
s (
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s and
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s) takes place in the seminiferous tubules of the testes. This process involves the production of several successive sperm cell precursors, starting with spermatogonia, which differentiate into spermatocytes. The spermatocytes then undergo meiosis, reducing their chromosome number by half, which produces spermatids. The spermatids then mature and, in animals, construct a tail, or flagellum, which gives rise to the mature, motile sperm cell. This whole process occurs constantly and takes around 3 months from start to finish. Sperm cells cannot divide and have a limited lifespan, but after fusion with
egg cell The egg cell, or ovum (plural ova), is the female reproductive cell, or gamete, in most anisogamous organisms (organisms that reproduce sexually with a larger, female gamete and a smaller, male one). The term is used when the female gamete is ...
s during
fertilisation Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proce ...
, a new organism begins developing, starting as a totipotent zygote. The human sperm cell is
haploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
, so that its 23 chromosomes can join the 23 chromosomes of the female egg to form a
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
cell with 46 paired chromosomes. In mammals, sperm is stored in the
epididymis The epididymis (; plural: epididymides or ) is a tube that connects a testicle to a vas deferens in the male reproductive system. It is a single, narrow, tightly-coiled tube in adult humans, in length. It serves as an interconnection between the ...
and is released from the penis during ejaculation in a fluid known as semen. The word ''sperm'' is derived from the Greek word σπέρμα, ''sperma'', meaning "seed".


Evolution

It is generally accepted that isogamy is the ancestor to sperm and eggs. However, there are no fossil records for the evolution of sperm and eggs from isogamy leading there to be a strong emphasis on mathematical models to understand the evolution of sperm. A widespread hypothesis states that sperm evolved rapidly, but there is no direct evidence that sperm evolved at a fast rate or before other male characteristics.


Sperm in animals


Function

The main sperm function is to reach the
ovum The egg cell, or ovum (plural ova), is the female reproductive cell, or gamete, in most anisogamous organisms (organisms that reproduce sexually with a larger, female gamete and a smaller, male one). The term is used when the female gamete is ...
and fuse with it to deliver two sub-cellular structures: (i) the male pronucleus that contains the genetic material and (ii) the
centriole In cell biology a centriole is a cylindrical organelle composed mainly of a protein called tubulin. Centrioles are found in most eukaryotic cells, but are not present in conifers (Pinophyta), flowering plants (angiosperms) and most fungi, and a ...
s that are structures that help organize the
microtubule Microtubules are polymers of tubulin that form part of the cytoskeleton and provide structure and shape to eukaryotic cells. Microtubules can be as long as 50 micrometres, as wide as 23 to 27  nm and have an inner diameter between 11 an ...
cytoskeleton.


Anatomy

The mammalian sperm cell can be divided in 2 parts: * Head: contains the nucleus with densely coiled chromatin fibers, surrounded anteriorly by a thin, flattened sac called the acrosome, which contains enzymes used for penetrating the female egg. It also contains vacuoles. * Tail: also called the flagellum, is the longest part and capable of wave-like motion that propels sperm for swimming and aids in the penetration of the egg. The tail was formerly thought to move symmetrically in a helical shape. The neck or connecting piece contains one typical centriole and one atypical centriole such as the
proximal centriole-like The proximal centriole-like or PCL is an atypical type of centriole found in the sperm cells of insects. The PCL name is due to some similarity to the Proximal centriole found in Vertebrates sperm and the hypothesis that the two structures are Homol ...
. The midpiece has a central filamentous core with many mitochondria spiralled around it, used for
ATP ATP may refer to: Companies and organizations * Association of Tennis Professionals, men's professional tennis governing body * American Technical Publishers, employee-owned publishing company * ', a Danish pension * Armenia Tree Project, non ...
production for the journey through the female
cervix The cervix or cervix uteri (Latin, 'neck of the uterus') is the lower part of the uterus (womb) in the human female reproductive system. The cervix is usually 2 to 3 cm long (~1 inch) and roughly cylindrical in shape, which changes during ...
, uterus, and uterine tubes. During fertilization, the sperm provides three essential parts to the oocyte: (1) a signalling or activating factor, which causes the
metabolically Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
dormant oocyte to activate; (2) the haploid paternal genome; (3) the centriole, which is responsible for forming the centrosome and
microtubule Microtubules are polymers of tubulin that form part of the cytoskeleton and provide structure and shape to eukaryotic cells. Microtubules can be as long as 50 micrometres, as wide as 23 to 27  nm and have an inner diameter between 11 an ...
system.


Origin

The spermatozoa of animals are produced through
spermatogenesis Spermatogenesis is the process by which haploid spermatozoa develop from germ cells in the seminiferous tubules of the testis. This process starts with the mitotic division of the stem cells located close to the basement membrane of the tubule ...
inside the male
gonad A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sper ...
s ( testicles) via
meiotic Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately res ...
division. The initial spermatozoon process takes around 70 days to complete. The process starts with the production of spermatogonia from germ cell precursors. These divide and differentiate into spermatocytes, which undergo meiosis to form spermatids. In the spermatid stage, the sperm develops the familiar tail. The next stage where it becomes fully mature takes around 60 days when it is called a
spermatozoan A spermatozoon (; also spelled spermatozoön; ; ) is a motile sperm cell (biology), cell, or moving form of the ploidy, haploid cell (biology), cell that is the male gamete. A spermatozoon Fertilization, joins an ovum to form a zygote. (A zygote ...
. Sperm cells are carried out of the male body in a fluid known as semen. Human sperm cells can survive within the female reproductive tract for more than 5 days post coitus. Semen is produced in the seminal vesicles, prostate gland and urethral glands. In 2016, scientists at
Nanjing Medical University Nanjing Medical University (NMU; ) is a university in Nanjing, Jiangsu Province, China. It was established in 1934 in Zhenjiang, but subsequently relocated to Nanjing in 1957. The university has two main campuses: Wutai and Jiangning, both of which ...
claimed they had produced cells resembling mouse spermatids from mouse embryonic stem cells artificially. They injected these spermatids into mouse eggs and produced pups.


Sperm quality

Sperm quantity and quality are the main parameters in semen quality, which is a measure of the ability of semen to accomplish fertilization. Thus, in humans, it is a measure of fertility in a man. The genetic quality of sperm, as well as its volume and motility, all typically decrease with age. DNA damages present in sperm cells in the period after meiosis but before fertilization may be repaired in the fertilized egg, but if not repaired, can have serious deleterious effects on fertility and the developing embryo. Human sperm cells are particularly vulnerable to free radical attack and the generation of oxidative DNA damage, such as that from 8-Oxo-2'-deoxyguanosine. The postmeiotic phase of mouse spermatogenesis is very sensitive to environmental genotoxic agents, because as male germ cells form mature sperm they progressively lose the ability to repair DNA damage. Irradiation of male mice during late spermatogenesis can induce damage that persists for at least 7 days in the fertilizing sperm cells, and disruption of maternal DNA double-strand break repair pathways increases sperm cell-derived chromosomal aberrations. Treatment of male mice with melphalan, a bifunctional alkylating agent frequently employed in chemotherapy, induces DNA lesions during meiosis that may persist in an unrepaired state as germ cells progress through DNA repair-competent phases of spermatogenic development. Such unrepaired DNA damages in sperm cells, after fertilization, can lead to offspring with various abnormalities.


Sperm size

Related to sperm quality is sperm size, at least in some animals. For instance, the sperm of some species of fruit fly ('' Drosophila'') are up to 5.8 cm long—about 20 times as long as the fly itself. Longer sperm cells are better than their shorter counterparts at displacing competitors from the female's seminal receptacle. The benefit to females is that only healthy males carry "good" genes that can produce long sperm in sufficient quantities to outcompete their competitors.


Market for human sperm

Some
sperm bank A sperm bank, semen bank, or cryobank is a facility or enterprise which purchases, stores and sells human semen. The semen is produced and sold by men who are known as sperm donors. The sperm is purchased by or for other persons for the purpose o ...
s hold up to of sperm. In addition to ejaculation, it is possible to extract sperm through testicular sperm extraction. On the global market, Denmark has a well-developed system of human sperm export. This success mainly comes from the reputation of Danish sperm donors for being of high quality and, in contrast with the law in the other Nordic countries, gives donors the choice of being either anonymous or non-anonymous to the receiving couple.Assisted Reproduction in the Nordic Countries
ncbio.org
Furthermore, Nordic sperm donors tend to be tall and highly educated
Posted Aug 13, 08 7:37 AM CDT in World, Science & Health
and have altruistic motives for their donations, partly due to the relatively low monetary compensation in Nordic countries. More than 50 countries worldwide are importers of Danish sperm, including Paraguay, Canada, Kenya, and Hong Kong. However, the Food and Drug Administration (FDA) of the US has banned import of any sperm, motivated by a risk of transmission of Creutzfeldt–Jakob disease, although such a risk is insignificant, since artificial insemination is very different from the route of transmission of Creutzfeldt–Jakob disease. The prevalence of Creutzfeldt–Jakob disease for donors is at most one in a million, and if the donor was a carrier, the infectious proteins would still have to cross the blood-testis barrier to make transmission possible.


History

Sperm were first observed in 1677 by
Antonie van Leeuwenhoek Antonie Philips van Leeuwenhoek ( ; ; 24 October 1632 – 26 August 1723) was a Dutch microbiologist and microscopist in the Golden Age of Dutch science and technology. A largely self-taught man in science, he is commonly known as " the ...
using a microscope. He described them as being animalcules (little animals), probably due to his belief in preformationism, which thought that each sperm contained a fully formed but small human.


Forensic analysis

Ejaculated fluids are detected by ultraviolet light, irrespective of the structure or colour of the surface. Sperm heads, e.g. from vaginal swabs, are still detected by
microscopy Microscopy is the technical field of using microscopes to view objects and areas of objects that cannot be seen with the naked eye (objects that are not within the resolution range of the normal eye). There are three well-known branches of micr ...
using the "Christmas Tree Stain" method, i.e., Kernechtrot-Picroindigocarmine (KPIC) staining.


Sperm in plants

Sperm cells in algal and many plant gametophytes are produced in male gametangia ( antheridia) via mitotic division. In flowering plants, sperm nuclei are produced inside
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
.


Motile sperm cells

Motile sperm cells typically move via flagella and require a water medium in order to swim toward the egg for fertilization. In animals most of the energy for sperm motility is derived from the metabolism of
fructose Fructose, or fruit sugar, is a Ketose, ketonic monosaccharide, simple sugar found in many plants, where it is often bonded to glucose to form the disaccharide sucrose. It is one of the three dietary monosaccharides, along with glucose and galacto ...
carried in the seminal fluid. This takes place in the
mitochondria A mitochondrion (; ) is an organelle found in the Cell (biology), cells of most Eukaryotes, such as animals, plants and Fungus, fungi. Mitochondria have a double lipid bilayer, membrane structure and use aerobic respiration to generate adenosi ...
located in the sperm's midpiece (at the base of the sperm head). These cells cannot swim backwards due to the nature of their propulsion. The uniflagellated sperm cells (with one flagellum) of animals are referred to as spermatozoa, and are known to vary in size. Motile sperm are also produced by many protists and the gametophytes of bryophytes, ferns and some gymnosperms such as cycads and ginkgo. The sperm cells are the only flagellated cells in the life cycle of these plants. In many ferns and lycophytes, cycads and ginkgo they are multi-flagellated (carrying more than one flagellum). In
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
s, the sperm cells are amoeboid and crawl, rather than swim, towards the egg cell.


Non-motile sperm cells

Non-motile sperm cells called spermatia lack flagella and therefore cannot swim. Spermatia are produced in a spermatangium. Because spermatia cannot swim, they depend on their environment to carry them to the egg cell. Some red algae, such as '' Polysiphonia'', produce non-motile spermatia that are spread by water currents after their release. The spermatia of rust fungi are covered with a sticky substance. They are produced in flask-shaped structures containing
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
, which attract flies that transfer the spermatia to nearby
hyphae A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...
for fertilization in a mechanism similar to insect pollination in flowering plants. Fungal spermatia (also called pycniospores, especially in the Uredinales) may be confused with
conidia A conidium ( ; ), sometimes termed an asexual chlamydospore or chlamydoconidium (), is an asexual, non-motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also called mitospores due to the ...
. Conidia are
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
s that germinate independently of fertilization, whereas spermatia are gametes that are required for fertilization. In some fungi, such as '' Neurospora crassa'', spermatia are identical to microconidia as they can perform both functions of fertilization as well as giving rise to new organisms without fertilization.


Sperm nuclei

In almost all embryophytes, including most gymnosperms and all
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s, the male gametophytes (
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
grains) are the primary mode of dispersal, for example via wind or insect pollination, eliminating the need for water to bridge the gap between male and female. Each pollen grain contains a spermatogenous (generative) cell. Once the pollen lands on the stigma of a receptive flower, it germinates and starts growing a
pollen tube A pollen tube is a tubular structure produced by the male gametophyte of seed plants when it germinates. Pollen tube elongation is an integral stage in the plant life cycle. The pollen tube acts as a conduit to transport the male gamete cells fro ...
through the carpel. Before the tube reaches the ovule, the nucleus of the generative cell in the pollen grain divides and gives rise to two sperm nuclei, which are then discharged through the tube into the ovule for fertilization. In some protists, fertilization also involves sperm nuclei, rather than cells, migrating toward the egg cell through a fertilization tube. Oomycetes form sperm nuclei in a syncytical antheridium surrounding the egg cells. The sperm nuclei reach the eggs through fertilization tubes, similar to the pollen tube mechanism in plants.


Sperm centrioles

Most sperm cells have centrioles in the sperm neck. Sperm of many animals has two typical centrioles, known as the proximal centriole and distal centriole. Some animals (including humans and bovines) have a single typical centriole, the proximal centriole, as well as a second centriole with atypical structure. Mice and rats have no recognizable sperm centrioles. The fruit fly '' Drosophila melanogaster'' has a single centriole and an atypical centriole named the proximal centriole-like.


Sperm tail formation

The sperm tail is a specialized type of cilium (aka flagella). In many animals the sperm tail is formed through the unique process of
cytosolic ciliogenesis Cytosolic ciliogenesis, otherwise cytoplasmic ciliogenesis, is a type of ciliogenesis where the cilium axoneme is formed in the cytoplasm or becomes exposed to the cytoplasm. Cytosolic ciliogenesis is divided into three types: Primary cytosolic ci ...
, in which all or part of the sperm tail's axoneme is formed in the cytoplasm or gets exposed to the cytoplasm.


See also

* Female sperm * Female sperm storage * Mendelian inheritance * Polyspermy * Sperm competition * Sperm granuloma * Sperm theft


Citations


General and cited sources

* Fawcett, D. W. (1981). "Sperm Flagellum". In: D. W. Fawcett
''The Cell'', 2nd ed
. Philadelphia: W. B. Saunders Company
pp. 604–640
. . . * Lehti, M. S. and A. Sironen (October 2017)
"Formation and function of sperm tail structures in association with sperm motility defects"
''Biol Reprod'' 97(4): 522–536. .


External links


Human Sperm Under a Microscope
{{Authority control Fertility Germ cells Reproductive system Mammal male reproductive system Semen Human male reproductive system Articles containing video clips Microscopic discoveries by Antonie van Leeuwenhoek