Spores, Bacterial
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In biology, a spore is a unit of sexual or
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 fu ...
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 Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
, fungi and
protozoa Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
. Bacterial spores are not part of a sexual cycle, but are resistant structures used for survival under unfavourable conditions. Myxozoan spores release amoeboid infectious germs ("amoebulae") into their hosts for parasitic infection, but also reproduce within the hosts through the pairing of two nuclei within the plasmodium, which develops from the amoebula. In plants, spores are usually
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 ...
and unicellular and are produced by meiosis in the
sporangium 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 ...
of 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 ...
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 pr ...
. Under favourable conditions the spore can develop into a new organism using mitotic division, producing a multicellular gametophyte, which eventually goes on to produce gametes. Two gametes fuse to form a zygote which develops into a new sporophyte. This cycle is known as
alternation of generations Alternation of generations (also known as metagenesis or heterogenesis) is the predominant type of Biological life cycle, life cycle in plants and algae. It consists of a Multicellular organism, multicellular haploid sexual phase, the gametophy ...
. The spores of seed plants are produced internally, and the megaspores (formed within the ovules) and the microspores are involved in the formation of more complex structures that form the dispersal units, the seeds and
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.


Definition

The term ''spore'' derives from the ancient Greek word σπορά ''spora'', meaning " seed, sowing", related to σπόρος ', "sowing", and σπείρειν ', "to sow". In common parlance, the difference between a "spore" and a " gamete" is that a spore will germinate and develop into a sporeling, while a gamete needs to combine with another gamete to form a zygote before developing further. The main difference between spores and seeds as dispersal units is that spores are unicellular, the first cell of a gametophyte, while seeds contain within them a developing embryo (the multicellular sporophyte of the next generation), produced by the fusion of the male gamete of the pollen tube with the female gamete formed by the megagametophyte within the ovule. Spores germinate to give rise to haploid gametophytes, while seeds germinate to give rise to diploid sporophytes.


Classification of spore-producing organisms


Plants

Vascular plant spores are always
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 ...
. Vascular plants are either homosporous (or isosporous) or heterosporous. Plants that are homosporous produce spores of the same size and type. Heterosporous plants, such as seed plants, spikemosses, quillworts, and ferns of the order Salviniales produce spores of two different sizes: the larger spore (megaspore) in effect functioning as a "female" spore and the smaller (microspore) functioning as a "male". Such plants typically give rise to the two kind of spores from within separate sporangia, either a megasporangium that produces megaspores or a microsporangium that produces microspores. In flowering plants, these sporangia occur within the carpel and anthers, respectively.


Fungi

Fungi commonly produce spores during sexual and asexual reproduction. Spores are usually
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 ...
and grow into mature haploid individuals through mitotic division of cells ( Urediniospores and Teliospores among rusts are dikaryotic). Dikaryotic cells result from the fusion of two haploid gamete cells. Among sporogenic dikaryotic cells, karyogamy (the fusion of the two haploid nuclei) occurs to produce a diploid cell. Diploid cells undergo meiosis to produce haploid spores.


Classification of spores

Spores can be classified in several ways:


By spore-producing structure


Fungi

In fungi and fungus-like organisms (e.g. Pseudofungi), spores are often classified by the structure in which meiosis and spore production occurs. Since fungi are often classified according to their spore-producing structures, these spores are often characteristic of a particular taxon of the fungi. *
Sporangiospore 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 cyc ...
s: spores produced by a
sporangium 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 ...
in many fungi such as zygomycetes. * Zygospores: spores produced by a zygosporangium, characteristic of zygomycetes. * Ascospores: spores produced by an
ascus An ascus (; ) is the sexual spore-bearing cell produced in ascomycete fungi. Each ascus usually contains eight ascospores (or octad), produced by meiosis followed, in most species, by a mitotic cell division. However, asci in some genera or s ...
, characteristic of
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. * Basidiospores: spores produced by a basidium, characteristic of basidiomycetes. * Aeciospores: spores produced by an
aecium An aecium (plural aecia) is a specialised reproductive structure found in some plant pathogenic rust fungi that produce aeciospores. Aecia may also be referred to as "cluster cups". The term aecidium (plural aecidia) is used interchangeably but i ...
in some fungi such as rusts or
smuts Smuts is an Afrikaans surname most commonly associated with Jan Smuts (1870–1950), a South African statesman, military leader and prime minister of the Union of South Africa. Other notable people with the surname include: * Barbara Smuts, Americ ...
. * Urediniospores: spores produced by a uredinium in some fungi such as rusts or
smuts Smuts is an Afrikaans surname most commonly associated with Jan Smuts (1870–1950), a South African statesman, military leader and prime minister of the Union of South Africa. Other notable people with the surname include: * Barbara Smuts, Americ ...
. * Teliospores: spores produced by a telium in some fungi such as rusts or
smuts Smuts is an Afrikaans surname most commonly associated with Jan Smuts (1870–1950), a South African statesman, military leader and prime minister of the Union of South Africa. Other notable people with the surname include: * Barbara Smuts, Americ ...
. * Oospores: spores produced by an
oogonium An oogonium (plural oogonia) is a small diploid cell which, upon maturation, forms a primordial follicle in a female fetus or the female (haploid or diploid) gametangium of certain thallophytes. In the mammalian fetus Oogonia are formed in larg ...
, characteristic of oomycetes.


Red algae

* Carpospores: spores produced by a carposporophyte, characteristic of
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 ...
. * Tetraspores: spores produced by a tetrasporophyte, characteristic of
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 ...
.


By function

* Chlamydospores: thick-walled resting spores of fungi produced to survive unfavorable conditions. * Parasitic fungal spores may be classified into internal spores, which germinate within the host, and external spores, also called environmental spores, released by the host to infest other hosts.


By origin during life cycle

* Meiospores: spores produced by meiosis; they are thus
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 ...
, and give rise to a haploid daughter cell(s) or a haploid individual. Examples are the precursor cells of gametophytes of seed plants found in flowers (
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) or cones ( gymnosperms), and the zoospores produced from meiosis in the sporophytes of algae such as '' Ulva''. ** Microspores: meiospores that give rise to a male gametophyte, (
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 ...
in seed plants). ** Megaspores (or macrospores): meiospores that give rise to a female gametophyte, (in seed plants the gametophyte forms within the ovule). * Mitospores (or conidia, conidiospores): spores produced by
mitosis In cell biology, mitosis () is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is mainta ...
; they are characteristic of
Ascomycetes 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 ...
. Fungi in which only mitospores are found are called "mitosporic fungi" or "anamorphic fungi", and are previously classified under the taxon Deuteromycota (See Teleomorph, anamorph and holomorph).


By mobility

Spores can be differentiated by whether they can move or not. * Zoospores: mobile spores that move by means of one or more
flagella A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...
, and can be found in some
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
and fungi. * Aplanospores: immobile spores that may nevertheless potentially grow flagella. * Autospores: immobile spores that cannot develop flagella. *
Ballistospore A ballistospore or ballistoconida is a spore that is discharged into the air from a living being, usually a species of fungus. With fungi, most types of basidiospores formed on basidia are discharged into the air from the tips of sterigmata. At leas ...
s: spores that are forcibly discharged or ejected from the fungal fruiting body as the result of an internal force, such as buildup of pressure. Most basidiospores are also ballistospores, and another notable example is spores of the genus '' Pilobolus''. * Statismospores: spores that are discharged from the fungal fruiting body as the result of an external force, such as raindrops or a passing animal. Examples are puffballs.


External anatomy

Under high
magnification Magnification is the process of enlarging the apparent size, not physical size, of something. This enlargement is quantified by a calculated number also called "magnification". When this number is less than one, it refers to a reduction in siz ...
, spores often have complex patterns or ornamentation on their exterior surfaces. A specialized terminology has been developed to describe features of such patterns. Some markings represent apertures, places where the tough outer coat of the spore can be penetrated when germination occurs. Spores can be categorized based on the position and number of these markings and apertures. Alete spores show no lines. In monolete spores, there is a single narrow line (laesura) on the spore. This indicates the mother spore split into four along a vertical axis. In trilete spores, each spore shows three narrow lines radiating from a center pole. This shows that four spores shared a common origin and were initially in contact with each other forming a tetrahedron. A wider aperture in the shape of a groove may be termed a colpus. The number of colpi distinguishes major groups of plants. Eudicots have tricolpate spores (i.e. spores with three colpi).


Spore tetrads and trilete spores

Envelope-enclosed spore tetrads are taken as the earliest evidence of plant life on land, dating from the mid-Ordovician (early Llanvirn, ~), a period from which no macrofossils have yet been recovered. Individual trilete spores resembling those of modern cryptogamic plants first appeared in the fossil record at the end of the Ordovician period.


Dispersal

In fungi, both asexual and sexual spores or sporangiospores of many fungal species are actively dispersed by forcible ejection from their reproductive structures. This ejection ensures exit of the spores from the reproductive structures as well as travelling through the air over long distances. Many fungi thereby possess specialized mechanical and physiological mechanisms as well as spore-surface structures, such as hydrophobins, for spore ejection. These mechanisms include, for example, forcible discharge of ascospores enabled by the structure of the ascus and accumulation of osmolytes in the fluids of the ascus that lead to explosive discharge of the ascospores into the air. The forcible discharge of single spores termed ''ballistospores'' involves formation of a small drop of water (
Buller's drop A basidium () is a microscopic sporangium (a spore-producing structure) found on the hymenophore of fruiting bodies of basidiomycete fungi which are also called tertiary mycelium, developed from secondary mycelium. Tertiary mycelium is highly ...
), which upon contact with the spore leads to its projectile release with an initial acceleration of more than 10,000 g. Other fungi rely on alternative mechanisms for spore release, such as external mechanical forces, exemplified by puffballs. Attracting insects, such as flies, to fruiting structures, by virtue of their having lively colours and a putrid odour, for dispersal of fungal spores is yet another strategy, most prominently used by the stinkhorns. In Common Smoothcap moss (''
Atrichum undulatum ''Atrichum undulatum'' is a species of mosses belonging to the family Polytrichaceae. It is native to Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Easter ...
''), the vibration of sporophyte has been shown to be an important mechanism for spore release. In the case of spore-shedding vascular plants such as ferns, wind distribution of very light spores provides great capacity for dispersal. Also, spores are less subject to animal predation than seeds because they contain almost no food reserve; however they are more subject to fungal and bacterial predation. Their chief advantage is that, of all forms of progeny, spores require the least energy and materials to produce. In the spikemoss ''
Selaginella lepidophylla ''Selaginella lepidophylla'' (syn. ''Lycopodium lepidophyllum'') is a species of desert plant in the spikemoss family (Selaginellaceae). Known as a "resurrection plant", ''S. lepidophylla'' is renowned for its ability to survive almost complete ...
'', dispersal is achieved in part by an unusual type of diaspore, a tumbleweed.


Gallery

File:Bartramia ithyphylla sporen.jpeg, Spores of the
moss Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hor ...
''Bartramia ithyphylla''. (microscopic view, 400x) File:Fernspore1.jpg, Dehisced fern sporangia. (microscopic view, no spores are visible) File:Botbrush1.jpg, Spores and elaters from a horsetail. ('' Equisetum'', microscopic view) File:Trilete spores.png, Fossil plant spores ('' Scylaspora'') from Silurian deposits of Sweden. File:Unknown fruit mold with spores methylene blue x2000.jpg, Fruit mold with spores and distinguishable cellular growth. (2000x) File:Reticularia olivacea-1.jpg, Spore clusters, formed inside sporangia of the slime mold ''
Reticularia olivacea Reticularia may refer to: * ''Reticularia'' (protist) Bull., 1788, a genus of protists * ''Reticularia'' Baumg., 1790, nom. illeg., synonym of ''Lobaria'' (Lobariaceae, Fungi) * ''Reticularia'' (brachiopod) M'Coy, 1844, a genus of Paleozoic Brach ...
'', from pine forests of eastern Ukraine. File:Tubifera dudkae-4.jpg, Internal surface of the peridium of the slime mold ''Tubifera dudkae'' with spores.


See also

* Aeroplankton *
Alternation of generations Alternation of generations (also known as metagenesis or heterogenesis) is the predominant type of Biological life cycle, life cycle in plants and algae. It consists of a Multicellular organism, multicellular haploid sexual phase, the gametophy ...
* Auxiliary cell *
Bioaerosol Bioaerosols (short for biological aerosols) are a subcategory of particles released from terrestrial and marine ecosystems into the atmosphere. They consist of both living and non-living components, such as fungi, pollen, bacteria and viruses. Comm ...
*
Cryptospores Cryptospores are fossilised primitive plant spores that first appear in the fossil record during the middle of the Ordovician period. Evidence that cryptospores derive from land plants Occurrence Cryptospores are generally found in non-marine ro ...
*
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., no ...
* Evolutionary history of plants * Fern *
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 pr ...


References

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