Sporogenesis is the production of spores in
biology
Biology is the scientific study of life and living organisms. It is a broad natural science that encompasses a wide range of fields and unifying principles that explain the structure, function, growth, History of life, origin, evolution, and ...
. The term is also used to refer to the process of
reproduction
Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. There are two forms of reproduction: Asexual reproduction, asexual and Sexual ...
via spores. Reproductive spores were found to be formed in
eukaryotic
The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...
organisms, such as
plant
Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they are predominantly Photosynthesis, photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with c ...
s,
algae
Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthesis, photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular ...
and
fungi
A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
, during their normal reproductive
life cycle. Dormant spores are formed, for example by certain fungi and algae, primarily in response to unfavorable growing conditions. Most eukaryotic spores are
haploid
Ploidy () is the number of complete sets of chromosomes in a cell (biology), cell, and hence the number of possible alleles for Autosome, autosomal and Pseudoautosomal region, pseudoautosomal genes. Here ''sets of chromosomes'' refers to the num ...
and form through cell division, though some types are
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. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...
s or
dikaryon
The dikaryon (karyogamy) is a cell nucleus feature that is unique to certain fungi. (The green alga '' Derbesia'' had been long considered an exception, until the heterokaryotic hypothesis was challenged by later studies.) Compatible cell-types c ...
s and form through cell fusion.
Reproduction via spores
Reproductive spores are generally the result of cell division, most commonly
meiosis
Meiosis () is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, the sperm or egg cells. It involves two rounds of division that ultimately result in four cells, each with only one c ...
(e.g. in plant
sporophyte
A sporophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the biological life cycle, life cycles of plants and algae. It is a diploid multicellular organism which produces asexual Spo ...
s). Sporic meiosis is needed to complete the sexual life cycle of the organisms using it.
In some cases, sporogenesis occurs via
mitosis
Mitosis () is a part of the cell cycle in eukaryote, eukaryotic cells in which replicated chromosomes are separated into two new Cell nucleus, nuclei. Cell division by mitosis is an equational division which gives rise to genetically identic ...
(e.g. in some fungi and algae). Mitotic sporogenesis is a form of
asexual reproduction
Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the f ...
. Examples are the
conidial fungi ''
Aspergillus
' () is a genus consisting of several hundred mold species found in various climates worldwide.
''Aspergillus'' was first catalogued in 1729 by the Italian priest and biologist Pier Antonio Micheli. Viewing the fungi under a microscope, Miche ...
'' and ''
Penicillium
''Penicillium'' () is a genus of Ascomycota, ascomycetous fungus, fungi that is part of the mycobiome of many species and is of major importance in the natural environment, in food spoilage, and in food and drug production.
Some members of th ...
'', for which mitospore formation appears to be the primary mode of reproduction. Other fungi, such as
ascomycete
Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The def ...
s, utilize both mitotic and meiotic spores. The red alga ''
Polysiphonia'' alternates between mitotic and meiotic sporogenesis and both processes are required to complete its complex reproductive life cycle.
In the case of dormant spores in eukaryotes, sporogenesis often occurs as a result of
fertilization
Fertilisation or fertilization (see American and British English spelling differences#-ise, -ize (-isation, -ization), spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give ...
or
karyogamy forming a diploid spore equivalent to a
zygote
A zygote (; , ) is a eukaryote, eukaryotic cell (biology), cell formed by a fertilization event between two gametes.
The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individ ...
. Therefore, zygospores are the result of
sexual reproduction
Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote tha ...
.
Reproduction via spores involves the spreading of the spores by water or air. Algae and some fungi (
chytrids) often use motile
zoospore
A zoospore is a motile asexual spore that uses a flagellum for locomotion in aqueous or moist environments. Also called a swarm spore, these spores are created by some protists, bacteria, and fungi to propagate themselves. Certain zoospores are ...
s that can swim to new locations before developing into sessile organisms. Airborne spores are obvious in fungi, for example when they are released from
puffballs. Other fungi have more active spore dispersal mechanisms. For example, the fungus ''
Pilobolus'' can shoot its sporangia towards light. Plant spores designed for
dispersal are also referred to as
diaspore
Diaspore ()also called diasporite, empholite, kayserite, or tanatariteis an aluminium hydroxide oxide mineral, α-AlO(OH), crystallizing in the orthorhombic system and isomorphous with goethite. It occurs sometimes as flattened crystals, but ...
s. Plant spores are most obvious in the reproduction of
fern
The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissue ...
s and
moss
Mosses are small, non-vascular plant, non-vascular flowerless plants in the taxonomic phylum, division Bryophyta (, ) ''sensu stricto''. Bryophyta (''sensu lato'', Wilhelm Philippe Schimper, Schimp. 1879) may also refer to the parent group bryo ...
es. However, they also exist in
flowering plant
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
s where they develop hidden inside the flower. For example, the
pollen grains of flowering plants develop out of microspores produced in the
anther
The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10
Morphology and terminology
A stamen typically consists of a stalk called the filament ...
s.
Reproductive spores grow into multicellular haploid individuals or
sporelings. In
heterosporous organisms, two types of spores exist: microspores give rise to males and megaspores to females. In
homosporous organisms, all spores look alike and grow into individuals carrying reproductive parts of both genders.
Formation of reproductive spores
Sporogenesis occurs in reproductive structures termed
sporangia
A sporangium (from Late Latin, ; : sporangia) is an enclosure in which spores are formed. It can be composed of a unicellular organism, single cell or can be multicellular organism, multicellular. Virtually all plants, fungus, fungi, and many ot ...
. The process involves sporogenous cells (sporocytes, also called spore mother cells) undergoing cell division to give rise to spores.
In meiotic sporogenesis, a diploid spore mother cell within the sporangium undergoes meiosis, producing a
tetrad of haploid spores. In organisms that are
heterosporous, two types of spores occur: Microsporangia produce
male
Male (Planet symbols, symbol: ♂) is the sex of an organism that produces the gamete (sex cell) known as sperm, which fuses with the larger female gamete, or Egg cell, ovum, in the process of fertilisation. A male organism cannot sexual repro ...
microspores, and megasporangia produce
female
An organism's sex is female ( symbol: ♀) if it produces the ovum (egg cell), the type of gamete (sex cell) that fuses with the male gamete (sperm cell) during sexual reproduction.
A female has larger gametes than a male. Females and ...
megaspores. In megasporogenesis, often three of the four spores degenerate after meiosis, whereas in microsporogenesis all four microspores survive.
In
gymnosperm
The gymnosperms ( ; ) are a group of woody, perennial Seed plant, seed-producing plants, typically lacking the protective outer covering which surrounds the seeds in flowering plants, that include Pinophyta, conifers, cycads, Ginkgo, and gnetoph ...
s, such as
conifer
Conifers () are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a sin ...
s, microspores are produced through meiosis from microsporocytes in
microstrobili or male cones. In
flowering plant
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
s, microspores are produced in the
anther
The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10
Morphology and terminology
A stamen typically consists of a stalk called the filament ...
s of flowers. Each anther contains four
pollen sacs, which contain the microsporocytes. After meiosis, each microspore undergoes mitotic cell division, giving rise to multicellular
pollen grains (six nuclei in gymnosperms, three nuclei in flowering plants).
Megasporogenesis occurs in
megastrobili in conifers (for example a pine cone) and inside the
ovule
In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the ''integument'', forming its outer layer, the ''nucellus'' (or remnant of the sporangium, megasporangium), ...
in the flowers of flowering plants. A megasporocyte inside a megasporangium or ovule undergoes meiosis, producing four megaspores. Only one is a functional megaspore whereas the others stay dysfunctional or degenerate. The megaspore undergoes several mitotic divisions to develop into a female
gametophyte
A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the se ...
(for example the seven-cell/eight-nuclei
embryo sac in flowering plants).
Mitospore formation
Some fungi and algae produce
mitospores through mitotic cell division within a sporangium. In fungi, such mitospores are referred to as
conidia
A conidium ( ; : conidia), sometimes termed an asexual chlamydospore or chlamydoconidium (: chlamydoconidia), is an asexual, non- motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also ...
.
Formation of dormant spores
Some algae, and fungi form
resting spores made to survive unfavorable conditions. Typically, changes in the environment from favorable to unfavorable growing conditions will trigger a switch from
asexual reproduction
Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the f ...
to
sexual reproduction
Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote tha ...
in these organisms. The resulting spores are protected through the formation of a thick
cell wall
A cell wall is a structural layer that surrounds some Cell type, cell types, found immediately outside the cell membrane. It can be tough, flexible, and sometimes rigid. Primarily, it provides the cell with structural support, shape, protection, ...
and can withstand harsh conditions such as drought or extreme temperatures. Examples are
chlamydospores,
teliospore
Teliospore (sometimes called teleutospore) is the thick-walled resting spore of some fungi (Rust (fungus), rusts and Smut (fungus), smuts), from which the basidium arises.
Development
They develop in ''telium, telia'' (sing. ''telium'' or ''telio ...
s,
zygospores, and
myxospores. Similar survival structures produced in some
bacteria
Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...
are known as
endospore
An endospore is a dormant, tough, and non-reproductive structure produced by some bacteria in the phylum Bacillota. The name "endospore" is suggestive of a spore or seed-like form (''endo'' means 'within'), but it is not a true spore (i.e., not ...
s.
Chlamydospore and teliospore formation
Chlamydospores are generally multicellular, asexual structures.
Teliospore
Teliospore (sometimes called teleutospore) is the thick-walled resting spore of some fungi (Rust (fungus), rusts and Smut (fungus), smuts), from which the basidium arises.
Development
They develop in ''telium, telia'' (sing. ''telium'' or ''telio ...
s are a form of chlamydospore produced through the fusion of cells or
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 o ...
where the
nuclei of the fused cells stay separate. These nuclei undergo
karyogamy and meiosis upon germination of the spore.
Zygospore, oospore and auxospore formation
Zygospores are formed in certain fungi (
zygomycota
Zygomycota, or zygote fungi, is a former phylum, division or phylum of the kingdom Fungi. The members are now part of two Phylum, phyla: the Mucoromycota and Zoopagomycotina, Zoopagomycota. Approximately 1060 species are known. They are mostly t ...
, for example ''
Rhizopus'') and some algae (for example ''
Chlamydomonas''). The zygospore forms through the
isogamic fusion of two cells (motile single cells in ''Chlamydomonas'') or sexual
conjugation
Conjugation or conjugate may refer to:
Linguistics
*Grammatical conjugation, the modification of a verb from its basic form
*Emotive conjugation or Russell's conjugation, the use of loaded language
Mathematics
*Complex conjugation, the change o ...
between two hyphae (in zygomycota).
Plasmogamy is followed by
karyogamy, therefore zygospores are diploid (
zygote
A zygote (; , ) is a eukaryote, eukaryotic cell (biology), cell formed by a fertilization event between two gametes.
The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individ ...
s). They will undergo zygotic meiosis upon germinating.
In
oomycetes, the zygote forms through the fertilization of an egg cell with a sperm nucleus and enters a resting stage as a diploid, thick-walled
oospore. The germinating oospore undergoes mitosis and gives rise to diploid hyphae which reproduce asexually via mitotic zoospores as long as conditions are favorable.
In
diatom
A diatom (Neo-Latin ''diatoma'') is any member of a large group comprising several Genus, genera of algae, specifically microalgae, found in the oceans, waterways and soils of the world. Living diatoms make up a significant portion of Earth's B ...
s, fertilization gives rise to a zygote termed
auxospore. Besides sexual reproduction and as a resting stage, the function of an auxospore is the restoration of the original cell size, as diatoms get progressively smaller during mitotic cell division. Auxospores divide by mitosis.
Endospore formation
The term sporogenesis can also refer to
endospore
An endospore is a dormant, tough, and non-reproductive structure produced by some bacteria in the phylum Bacillota. The name "endospore" is suggestive of a spore or seed-like form (''endo'' means 'within'), but it is not a true spore (i.e., not ...
formation in
bacteria
Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...
, which allows the cells to survive unfavorable conditions. Endospores are not reproductive structures and their formation does not require cell fusion or division. Instead, they form through the production of an encapsulating spore coat within the spore-forming cell.
Parts of the spore
There are many parts of the spore 'plant'. The structure enclosing a group of spores is called a
sporangium
A sporangium (from Late Latin, ; : sporangia) is an enclosure in which spores are formed. It can be composed of a unicellular organism, single cell or can be multicellular organism, multicellular. Virtually all plants, fungus, fungi, and many ot ...
.
Bibliography
*S.S. Mader (2007): ''Biology'', 9th edition, McGraw Hill Companies, New York,
*P.H. Raven, R.F. Evert, S.E. Eichhorn (2005): ''Biology of Plants'', 7th Edition, W.H. Freeman and Company Publishers, New York, {{ISBN, 0-7167-1007-2
Reproduction
Reproductive system
Plant reproduction