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Sporogenesis is the production of spores in
biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary ...
. The term is also used to refer to the process of reproduction via spores. Reproductive spores were found to be formed in eukaryotic organisms, such as plants,
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthesis, photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from u ...
and fungi, during their normal reproductive
life cycle Life cycle, life-cycle, or lifecycle may refer to: Science and academia *Biological life cycle, the sequence of life stages that an organism undergoes from birth to reproduction ending with the production of the offspring *Life-cycle hypothesis, ...
. Dormant spores are formed, for example by certain fungi and algae, primarily in response to unfavorable growing conditions. Most eukaryotic spores are haploid and form through cell division, though some types are diploid or dikaryons and form through cell fusion.


Reproduction via spores

Reproductive spores are generally the result of cell division, most commonly meiosis (e.g. in plant
sporophyte A sporophyte () is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase. Life cycle The sporophyte develops from the zygote p ...
s). Sporic meiosis is needed to complete the sexual life cycle of the organisms using it. In some cases, sporogenesis occurs via mitosis (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, Mich ...
'' and ''
Penicillium ''Penicillium'' () is a genus of ascomycetous 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 the genus produce pe ...
'', 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 defi ...
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 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 ...
or karyogamy forming a diploid spore equivalent to a zygote. Therefore, zygospores are the result of sexual reproduction. Reproduction via spores involves the spreading of the spores by water or air. Algae and some fungi ( chytrids) often use motile zoospores that can swim to new locations before developing into sessile organisms. Airborne spores are obvious in fungi, for example when they are released from
puffball Puffballs are a type of fungus featuring a ball-shaped fruit body that bursts on impact, releasing a cloud of dust-like spores when mature. Puffballs belong to the division Basidiomycota and encompass several genera, including '' Calvatia'', '' ...
s. Other fungi have more active spore dispersal mechanisms. For example, the fungus ''
Pilobolus ''Pilobolus'' is a genus of fungi that commonly grows on herbivore dung. Life cycle The life cycle of ''Pilobolus'' begins with a black sporangium that has been discharged onto a plant substrate such as grass. A herbivorous animal such as a ...
'' can shoot its sporangia towards light. Plant spores designed for dispersal are also referred to as diaspores. Plant spores are most obvious in the reproduction of ferns and mosses. However, they also exist in flowering plants where they develop hidden inside the flower. For example, the
pollen grain 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 gametoph ...
s of flowering plants develop out of microspores produced in the
anther The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filam ...
s. Reproductive spores grow into multicellular haploid individuals or
sporeling A sporeling is a young plant or fungus produced by a germinated spore, similar to a seedling derived from a germinated seed. They occur in algae, fungi, lichens, bryophytes and seedless vascular plants. Sporeling development Most spores germin ...
s. In heterosporous organisms, two types of spores exist: microspores give rise to males and megaspores to females. In
homosporous 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, ...
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, ) is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other lineages form sporangia at some point in their life cy ...
. 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 Tetrad ('group of 4') or tetrade may refer to: * Tetrad (area), an area 2 km x 2 km square * Tetrad (astronomy), four total lunar eclipses within two years * Tetrad (chromosomal formation) * Tetrad (general relativity), or frame field ** Tetra ...
of haploid spores. In organisms that are heterosporous, two types of spores occur: Microsporangia produce male microspores, and megasporangia produce female megaspores. In megasporogenesis, often three of the four spores degenerate after meiosis, whereas in microsporogenesis all four microspores survive. In gymnosperms, such as conifers, microspores are produced through meiosis from microsporocytes in microstrobili or male cones. In flowering plants, microspores are produced in the
anther The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filam ...
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 grain 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 gametoph ...
s (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 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 (for example the seven-cell/eight-nuclei
embryo sac 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 megasporangium), and the fem ...
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 ( ; ), 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 t ...
.


Formation of dormant spores

Some algae, and fungi form
resting spore A resting spore is a resistant cell, used to survive adverse environmental conditions. Resting spore is a term commonly applied to both diatoms and fungi. In fungi A resting spore can be a spore created by fungi which is thickly encysted (has a t ...
s 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 in these organisms. The resulting spores are protected through the formation of a thick
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering me ...
and can withstand harsh conditions such as drought or extreme temperatures. Examples are
chlamydospore A chlamydospore is the thick-walled large resting spore of several kinds of fungi, including Ascomycota such as '' Candida'', Basidiomycota such as '' Panus'', and various Mortierellales species. It is the life-stage which survives in unfavourable ...
s,
teliospore Teliospore (sometimes called teleutospore) is the thick-walled resting spore of some fungi ( rusts and smuts), from which the basidium arises. Development They develop in '' telia'' (sing. ''telium'' or ''teliosorus''). The telial host is the ...
s,
zygospore A zygospore is a diploid reproductive stage in the life cycle of many fungi and protists. Zygospores are created by the nuclear fusion of haploid cells. In fungi, zygospores are formed in zygosporangia after the fusion of specialized budding struc ...
s, and myxospores. Similar survival structures produced in some bacteria are known as endospores.


Chlamydospore and teliospore formation

Chlamydospore A chlamydospore is the thick-walled large resting spore of several kinds of fungi, including Ascomycota such as '' Candida'', Basidiomycota such as '' Panus'', and various Mortierellales species. It is the life-stage which survives in unfavourable ...
s are generally multicellular, asexual structures.
Teliospore Teliospore (sometimes called teleutospore) is the thick-walled resting spore of some fungi ( rusts and smuts), from which the basidium arises. Development They develop in '' telia'' (sing. ''telium'' or ''teliosorus''). The telial host is the ...
s are a form of chlamydospore produced through the fusion of cells or hyphae 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

Zygospore A zygospore is a diploid reproductive stage in the life cycle of many fungi and protists. Zygospores are created by the nuclear fusion of haploid cells. In fungi, zygospores are formed in zygosporangia after the fusion of specialized budding struc ...
s are formed in certain fungi (
zygomycota Zygomycota, or zygote fungi, is a former division or phylum of the kingdom Fungi. The members are now part of two phyla: the Mucoromycota and Zoopagomycota. Approximately 1060 species are known. They are mostly terrestrial in habitat, living i ...
, for example ''
Rhizopus ''Rhizopus'' is a genus of common saprophytic fungi on plants and specialized parasites on animals. They are found in a wide variety of organic substances, including "mature fruits and vegetables", jellies, syrups, leather, bread, peanuts, and t ...
'') and some algae (for example ''
Chlamydomonas ''Chlamydomonas'' is a genus of green algae consisting of about 150 speciesSmith, G.M. 1955 ''Cryptogamic Botany Volume 1. Algae and Fungi'' McGraw-Hill Book Company Inc of unicellular flagellates, found in stagnant water and on damp soil, i ...
''). 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 ...
between two hyphae (in zygomycota).
Plasmogamy Plasmogamy is a stage in the sexual reproduction of fungi, in which the protoplasm of two parent cells (usually from the mycelia) fuse without the fusion of nuclei, effectively bringing two haploid nuclei close together in the same cell. This sta ...
is followed by karyogamy, therefore zygospores are diploid ( zygotes). They will undergo zygotic meiosis upon germinating. In
oomycete Oomycota forms a distinct phylogenetic lineage of fungus-like eukaryotic microorganisms, called oomycetes (). They are filamentous and heterotrophic, and can reproduce both sexually and asexually. Sexual reproduction of an oospore is the resul ...
s, 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''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising se ...
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 formation in bacteria, 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 case that holds all the spores is called a sporangium.


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