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Spores produced in a sporic life cycle. Sporic meiosis.svg
Spores produced in a sporic life cycle.
Fresh snow partially covers rough-stalked feather-moss (Brachythecium rutabulum), growing on a thinned hybrid black poplar (Populus x canadensis). The last stage of the moss lifecycle is shown, where the sporophytes are visible before dispersion of their spores: the calyptra (1) is still attached to the capsule (3). The tops of the gametophytes (2) can be discerned as well. Inset shows the surrounding, black poplars growing on sandy loam on the bank of a kolk, with the detail area marked. Brachythecium rutabulum on Populus x canadensis.jpg
Fresh snow partially covers rough-stalked feather-moss (Brachythecium rutabulum), growing on a thinned hybrid black poplar (Populus x canadensis). The last stage of the moss lifecycle is shown, where the sporophytes are visible before dispersion of their spores: the calyptra (1) is still attached to the capsule (3). The tops of the gametophytes (2) can be discerned as well. Inset shows the surrounding, black poplars growing on sandy loam on the bank of a kolk, with the detail area marked.

In biology, a spore is a unit of sexual (in fungi) or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. [1] Spores form part of the life cycles of many plants, algae, fungi and protozoa. [2] They were thought to have appeared as early as the mid-late Ordovician period as an adaptation of early land plants. [3]


Bacterial spores are not part of a sexual cycle, but are resistant structures used for survival under unfavourable conditions. [4] 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. [5]

In plants, spores are usually haploid and unicellular and are produced by meiosis in the sporangium of a diploid sporophyte. In some rare cases, diploid spore is also produced in some algae, or fungi. [6] 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.

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 grains.


The term spore derives from the ancient Greek word σπορά spora, meaning "seed, sowing", related to σπόρος sporos, "sowing", and σπείρειν speirein, "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


Vascular plant spores are always haploid. 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 commonly produce spores during sexual and asexual reproduction. Spores are usually haploid 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 such as by their spore producing structure, function, origin during life cycle, and mobility.

Below is a table listing the mode of classification, name, identifying characteristic, examples, and images of different spore species.

Mode of ClassificationNameIdentifiying CharacteristicExample Spore Containing OrganismImage
Spore Producing Structure Sporangiospore Produced by sporangium Zygomycetes
Sporangium of Fungi Sporangium of fungi.jpg
Sporangium of Fungi
Zygospores Produced by zygosporangium Zygomycetes
Zygospores on Rhizopus Rhizopus zygospores.jpg
Zygospores on Rhizopus
Ascospores Produced by ascus Ascomycetes
Ascospores of Didymella Rabiei Bitunicate ascus and ascospores of Didymella rabiei.png
Ascospores of Didymella Rabiei
Basidiospores Produced by basidium Basidiomycetes
Typical reproductive structure of a basidiomycete, including the basidiospore and basidium Basidium schematic.svg
Typical reproductive structure of a basidiomycete, including the basidiospore and basidium
Aecispores Produced by aecium Rusts and Smuts
Aceium on foilage Aecium sp.jpg
Aceium on foilage
Urediniospores Produced by urediniumRusts and Smuts
Uredinospores Puccinia thaliae urediniospores.jpg
Teliospores Produced by teilum Rusts and Smuts
Microscopic image of teliospores Puccinia helianthi (teliospores).jpg
Microscopic image of teliospores
Oospores Produced by oogonium Oomycetes
Oospores of Phytophthora agathidicida 10554 oospore.jpg
Oospores of Phytophthora agathidicida
Carpospores Produced by carposophorophyte Red Algae
Light microscopy of Polysiphonia showing a carpospores and carposporophyte inside Rhod1004.jpg
Light microscopy of Polysiphonia showing a carpospores and carposporophyte inside
Tetraspores Produced by tetrasphorophyte Red Algae
Tetraspores of Polysiphonia Polysiphonia tetraspores WM2.jpg
Tetraspores of Polysiphonia
Function Chalmydospore Thick-walled resting spores of fungi produced to survive in unfavorable conditions Asomycota
Pseudohyphae, chlamydospores and blastospores of Candida yeast. Candida pseudohyphae, chlamydospores, blastospores.png
Pseudohyphae, chlamydospores and blastospores of Candida yeast.
Parasitic Fungal SporeInternal SporesGerminate within a host
A parasitic pink fungi on a Lichen tree Pink fungal parasite on lichen - geograph.org.uk - 1040204.jpg
A parasitic pink fungi on a Lichen tree
External (Environmental) sporesSpores released by the host to infest other hosts [7]
Origin During Life CycleMeiospores Microspores Produced sexually through meiosis, and give rise to a male gametophyte Pollen in seed plants
In plants, microspores, and in some cases megaspores, are formed from all four products of meiosis. Microspore-formation.svg
In plants, microspores, and in some cases megaspores, are formed from all four products of meiosis.
Megaspores (macrospores)Produced sexually through meiosis, and give rise to a female gametophyte Ovule in seed plants
In contrast, in many seed plants and heterosporous ferns, only a single product of meiosis will become a megaspore (macrospore), with the rest degenerating. Macrospore-formation.svg
In contrast, in many seed plants and heterosporous ferns, only a single product of meiosis will become a megaspore (macrospore), with the rest degenerating.
Mitospores Produced asexually though mitosis Ascomycetes
Ascomycete containing mitospores Morchella conica 1 beentree.jpg
Ascomycete containing mitospores
Mobility Zoospores Mobile through flagella Some algae and fungi
Microscopic image of a Zoospore Zoospores - Reproductive Structure of the Phytophthora.png
Microscopic image of a Zoospore
AplanosporesImmobile, however still produce flagella
Autospores Immobile spores that do not produce flagella
Autospores of a strain of Jenufa aeroterrestrica Jenufa SAG 2383.jpg
Autospores of a strain of Jenufa aeroterrestrica
Ballistospores Forcibly discharged from the fungal fruiting body due to internal force (such as built up pressure)Basidiospores and/or part of the genus Pilobus
Ballistospore mechanism of dispersal from fungi Abb2.5 Fungi Basidiomycota basidium basidiospore ballistospore Bullers drop catapult mechanism 2021 (M. Piepenbring).png
Ballistospore mechanism of dispersal from fungi
StratismosporesForcibly discharged from the fungal fruting body due to external force (such as raindrops or passing animals) Puffballs
Puff Balls containing Stratismospores Puff Balls - geograph.org.uk - 6255537.jpg
Puff Balls containing Stratismospores

External anatomy

Fossil trilete spores (blue) and a spore tetrad (green) of Late Silurian origin Trilete spores.png
Fossil trilete spores (blue) and a spore tetrad (green) of Late Silurian origin
Tricolpate pollen of Ricinus Pollen Ricinus communis sanguineus.jpg
Tricolpate pollen of Ricinus

Under high magnification, 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. [8] Indicating the prior contact of two spores that eventually separated. [3] In trilete spores, each spore shows three narrow lines radiating from a center pole. [8] This shows that four spores shared a common origin and were initially in contact with each other forming a tetrahedron. [3] A wider aperture in the shape of a groove may be termed a colpus. [8] The number of colpi distinguishes major groups of plants. Eudicots have tricolpate spores (i.e. spores with three colpi). [9]

Spore tetrads and trilete spores

Envelope-enclosed spore tetrads are taken as the earliest evidence of plant life on land, [10] dating from the mid-Ordovician (early Llanvirn, ~ 470  million years ago), a period from which no macrofossils have yet been recovered. [11] Individual trilete spores resembling those of modern cryptogamic plants first appeared in the fossil record at the end of the Ordovician period. [12]


Spores being ejected by fungi.

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. [13]

The forcible discharge of single spores termed ballistospores involves formation of a small drop of water (Buller's drop), which upon contact with the spore leads to its projectile release with an initial acceleration of more than 10,000 g. [14] 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 ), the vibration of sporophyte has been shown to be an important mechanism for spore release. [15]

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 , dispersal is achieved in part by an unusual type of diaspore, a tumbleweed. [16]


Spores have been found in microfossils dating back to the mid-late Ordovician period. [17] Two hypothesized initial functions of spores relate to whether they appeared before or after land plants. The heavily studied hypothesis is that spores were an adaptation of early land plant species, such as embryophytes, that allowed for plants to easily disperse while adapting to their non-aquatic environment. [17] [18] This is particularly supported by the observation of a thick spore wall in cryptospores. These spore walls would have protected potential offspring from novel weather elements. [17] The second more recent hypothesis is that spores were an early predecessor of land plants and formed during errors in the meiosis of algae, a hypothesized early ancestor of land plants. [19]

Whether spores arose before or after land plants, their contributions to topics in fields like paleontology and plant phylogenetics have been useful . [19] The spores found in microfossils, also known as cryptospores, are well preserved due to the fixed material they are in as well as how abundant and widespread they were during their respective time periods. These microfossils are especially helpful when studying the early periods of earth as macrofossils such as plants are not common nor well preserved. [17] Both cryptospores and modern spores have diverse morphology that indicate possible environmental conditions of earlier periods of Earth and evolutionary relationships of plant species. [17] [19] [18]

See also

Related Research Articles

<span class="mw-page-title-main">Gametophyte</span> Haploid stage in the life cycle of plants and algae

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 sexual phase in the life cycle of plants and algae. It develops sex organs that produce gametes, haploid sex cells that participate in fertilization to form a diploid zygote which has a double set of chromosomes. Cell division of the zygote results in a new diploid multicellular organism, the second stage in the life cycle known as the sporophyte. The sporophyte can produce haploid spores by meiosis that on germination produce a new generation of gametophytes.

<span class="mw-page-title-main">Sporangium</span> Enclosure in which spores are formed

A sporangium ; pl.: sporangia) 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 cycle. Sporangia can produce spores by mitosis, but in nearly all land plants and many fungi, sporangia are the site of meiosis and produce genetically distinct haploid spores.

<span class="mw-page-title-main">Alternation of generations</span> Reproductive cycle of plants and algae

Alternation of generations is the predominant type of life cycle in plants and algae. In plants both phases are multicellular: the haploid sexual phase – the gametophyte – alternates with a diploid asexual phase – the sporophyte.

<i>Isoetes</i> Genus of vascular plants in the family Isoetaceae

Isoetes, commonly known as the quillworts, is a genus of lycopod. It is the only living genus in the family Isoetaceae and order Isoetales. There are currently 192 recognized species, with a cosmopolitan distribution mostly in aquatic habitats but with the individual species often scarce to rare. Some botanists split the genus, separating two South American species into the genus Stylites, although molecular data place these species among other species of Isoetes, so that Stylites does not warrant taxonomic recognition. Species virtually identical to modern quillworts have existed since the Jurassic epoch, though the timing of the origin of modern Isoetes is subject to considerable uncertainty.

<span class="mw-page-title-main">Bryophyte</span> Terrestrial plants that lack vascular tissue

Bryophytes are a group of land plants, sometimes treated as a taxonomic division, that contains three groups of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. In the strict sense, Bryophyta consists of the mosses only. Bryophytes are characteristically limited in size and prefer moist habitats although they can survive in drier environments. The bryophytes consist of about 20,000 plant species. Bryophytes produce enclosed reproductive structures, but they do not produce flowers or seeds. They reproduce sexually by spores and asexually by fragmentation or the production of gemmae. Though bryophytes were considered a paraphyletic group in recent years, almost all of the most recent phylogenetic evidence supports the monophyly of this group, as originally classified by Wilhelm Schimper in 1879. The term bryophyte comes from Ancient Greek βρύον (brúon) 'tree moss, liverwort', and φυτόν (phutón) 'plant'.

<span class="mw-page-title-main">Gametogenesis</span> Biological process

Gametogenesis is a biological process by which diploid or haploid precursor cells undergo cell division and differentiation to form mature haploid gametes. Depending on the biological life cycle of the organism, gametogenesis occurs by meiotic division of diploid gametocytes into various gametes, or by mitosis. For example, plants produce gametes through mitosis in gametophytes. The gametophytes grow from haploid spores after sporic meiosis. The existence of a multicellular, haploid phase in the life cycle between meiosis and gametogenesis is also referred to as alternation of generations.

<span class="mw-page-title-main">Embryophyte</span> Subclade of green plants, also known as land plants

The embryophytes are a clade of plants, also known as Embryophyta or land plants. They are the most familiar group of photoautotrophs that make up the vegetation on Earth's dry lands and wetlands. Embryophytes have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of freshwater charophyte green algae as a sister taxon of Charophyceae, Coleochaetophyceae and Zygnematophyceae. Embryophytes consist of the bryophytes and the polysporangiophytes. Living embryophytes include hornworts, liverworts, mosses, lycophytes, ferns, gymnosperms and angiosperms. Embryophytes have diplobiontic life cycles.

<span class="mw-page-title-main">Biological life cycle</span> Series of stages of an organism

In biology, a biological life cycle is a series of stages of the life of an organism, that begins as a zygote, often in an egg, and concludes as an adult that reproduces, producing an offspring in the form of a new zygote which then itself goes through the same series of stages, the process repeating in a cyclic fashion.

<span class="mw-page-title-main">Sporophyte</span> Diploid multicellular stage in the life cycle of a plant or alga

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.

Dioecy is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly or indirectly. Dioecious reproduction is biparental reproduction. Dioecy has costs, since only the female part of the population directly produces offspring. It is one method for excluding self-fertilization and promoting allogamy (outcrossing), and thus tends to reduce the expression of recessive deleterious mutations present in a population. Plants have several other methods of preventing self-fertilization including, for example, dichogamy, herkogamy, and self-incompatibility.

Microsporangia are sporangia that produce microspores that give rise to male gametophytes when they germinate. Microsporangia occur in all vascular plants that have heterosporic life cycles, such as seed plants, spike mosses and the aquatic fern genus not species Azolla. In gymnosperms and angiosperm anthers, the microsporangia produce microsporocytes, the microspore mother cells, which then produce four microspores through the process of meiosis. Microsporocytes are produced in the microsporangia of gymnosperm cones and the anthers of angiosperms. They are diploid microspore mother-cells, which then produce four haploid microspores through the process of meiosis. These become pollen grains, within which the microspores divide twice by mitosis to produce a very simple gametophyte.

<span class="mw-page-title-main">Double fertilization</span> Complex fertilization mechanism of flowering plants

Double fertilization or Double fertilisation is a complex fertilization mechanism of flowering plants (angiosperms). This process involves the joining of a female gametophyte with two male gametes (sperm). It begins when a pollen grain adheres to the stigma of the carpel, the female reproductive structure of a flower. The pollen grain then takes in moisture and begins to germinate, forming a pollen tube that extends down toward the ovary through the style. The tip of the pollen tube then enters the ovary and penetrates through the micropyle opening in the ovule. The pollen tube proceeds to release the two sperm in the embryo sacs.

<span class="mw-page-title-main">Microspore</span> Small land plant spores that develop into male gametophytes

Microspores are land plant spores that develop into male gametophytes, whereas megaspores develop into female gametophytes. The male gametophyte gives rise to sperm cells, which are used for fertilization of an egg cell to form a zygote. Megaspores are structures that are part of the alternation of generations in many seedless vascular cryptogams, all gymnosperms and all angiosperms. Plants with heterosporous life cycles using microspores and megaspores arose independently in several plant groups during the Devonian period. Microspores are haploid, and are produced from diploid microsporocytes by meiosis.

Plant reproduction is the production of new offspring in plants, which can be accomplished by sexual or asexual reproduction. Sexual reproduction produces offspring by the fusion of gametes, resulting in offspring genetically different from either parent. Asexual reproduction produces new individuals without the fusion of gametes, resulting in clonal plants that are genetically identical to the parent plant and each other, unless mutations occur.

Sporogenesis is the production of spores in biology. 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 and fungi, 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 and form through cell division, though some types are diploid sor dikaryons and form through cell fusion.we can also say this type of reproduction as single pollination

In botany, a zoid or zoïd is a reproductive cell that possesses one or more flagella, and is capable of independent movement. Zoid can refer to either an asexually reproductive spore or a sexually reproductive gamete. In sexually reproductive gametes, zoids can be either male or female depending on the species. For example, some brown alga (Phaeophyceae) reproduce by producing multi-flagellated male and female gametes that recombine to form the diploid sporangia. Zoids are primarily found in some protists, diatoms, green alga, brown alga, non-vascular plants, and a few vascular plants. The most common classification group that produces zoids is the heterokonts or stramenopiles. These include green alga, brown alga, oomycetes, and some protists. The term is generally not used to describe motile, flagellated sperm found in animals. Zoid is also commonly confused for zooid which is a single organism that is part of a colonial animal.

<span class="mw-page-title-main">Embryonic sac</span>

A megaspore mother cell, or megasporocyte, is a diploid cell in plants in which meiosis will occur, resulting in the production of four haploid megaspores. At least one of the spores develop into haploid female gametophytes (megagametophytes). The megaspore mother cell arises within the megasporangium tissue.

<span class="mw-page-title-main">Heterospory</span> The production of spores of two different sizes and sexes by several groups of land plants

Heterospory is the production of spores of two different sizes and sexes by the sporophytes of land plants. The smaller of these, the microspore, is male and the larger megaspore is female. Heterospory evolved during the Devonian period from isospory independently in several plant groups: the clubmosses, the ferns including the arborescent horsetails, and progymnosperms. This occurred as part of the process of evolution of the timing of sex differentiation.

<span class="mw-page-title-main">Sexual reproduction</span> Biological process

Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete with a single set of chromosomes combines with another gamete to produce a zygote that develops into an organism composed of cells with two sets of chromosomes (diploid). This is typical in animals, though the number of chromosome sets and how that number changes in sexual reproduction varies, especially among plants, fungi, and other eukaryotes.

<i>Asterotheca</i> Genus of plants

Asterotheca is a genus of seedless, spore-bearing, vascularized ferns dating from the Carboniferous of the Paleozoic to the Triassic of the Mesozoic.


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