thumb|200px|''Equisetum arvense'' [[strobilus">Image:Equisetum arvense sporangia.jpg|thumb|200px|''Equisetum arvense'' [[strobilus
cut open to reveal sporangia
A sporangium (pl., sporangia) (modern Latin, from [[Ancient Greek language|Greek]] ''σπόρος ()'' ‘spore’ + ''ἀγγεῖον ()'' ‘vessel’) is an enclosure in which [[spores]] are formed. It can be composed of a single cell or can be multicellular. All [[plant]]s, 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.
In some phyla of fungi, the sporangium plays a role in asexual reproduction
, and may play an indirect role in sexual reproduction
. The sporangium forms on the sporangiophore and contains haploid nuclei
are formed in the sporangiophore by encasing each haploid nucleus and cytoplasm in a tough outer membrane
. During asexual reproduction
, these spores
are dispersed via wind and germinate
into haploid hypha
Although sexual reproduction
in fungi varies between phyla, for some fungi the sporangium plays an indirect role in sexual reproduction. For Zygomycota, sexual reproduction occurs when the haploid hyphae from two individuals join to form a zygosporangium in response to unfavorable conditions. The haploid nuclei within the zygosporangium then fuse into diploid
When conditions improve the zygosporangium germinates, undergoes meiosis and produces a sporangium, which releases spores.
In mosses, liverworts and hornworts, an unbranched sporophyte
produces a single sporangium, which may be quite complex morphologically. Most non-vascular plants, as well as many lycophytes and most ferns, are homosporous
(only one kind of spore is produced). Some bryophytes, most lycophytes, and some ferns are heterosporous
(two kinds of spores are produced). These plants produce microspores and megaspores, which give rise to gametophytes that are functionally male or female, respectively. In some cases, both kinds of spores are produced in the same sporangium, and may even develop together as part of a spore tetrad. However, in most heterosporous plants there are two kinds of sporangia, termed microsporangia
and megasporangia. A few ferns (Salviniaceae
) and some lycophytes (the genera ''Selaginella
'' and ''Isoetes
'' and the extinct lepidodendrids) are heterosporous with two kinds of sporangia, as are all the seed plants.
Sporangia can be terminal (on the tips) or lateral (placed along the side) of stems or associated with leaves. In fern
s, sporangia are typically found on the abaxial
surface (underside) of the leaf and are densely aggregated into clusters called sori
. Sori may be covered by a structure called an indusium. Some ferns have their sporangia scattered along reduced leaf segments or along (or just in from) the margin of the leaf.
Lycophytes, in contrast, bear their sporangia on the adaxial
surface (the upper side) of leaves or laterally on stems. Leaves that bear sporangia are called sporophyll
s. If the plant is heterosporous, the sporangia-bearing leaves are distinguished as either microsporophylls or megasporophylls. In seed plants, sporangia are typically located within strobili
s form their microsporangia on microsporophylls which are aggregated into strobili. Megasporangia are formed into ovules, which are borne on megasporophylls, which are aggregated into strobili on separate plants (all cycads are dioecious). Conifer
s typically bear their microsporangia on microsporophylls aggregated into papery pollen strobili, and the ovules, are located on modified stem axes forming compound ovuliferous cone scales. Flowering plants contain microsporangia in the anthers of stamen
s (typically four microsporangia per anther) and megasporangia inside ovule
s inside ovaries. In all seed plants, spores are produced by meiosis and develop into gametophytes while still inside the sporangium. The microspores become microgametophytes (pollen). The megaspores become megagametophytes (embryo sacs).
Eusporangia and leptosporangia
Categorized based on developmental sequence, ''eusporangia'' and ''leptosporangia'' are differentiated in the vascular plants.
* In a leptosporangium, found only in ferns, development involves a single initial cell that becomes the stalk, wall, and spores within the sporangium. There are around 64 spores in a leptosporangium.
* In a eusporangium, characteristic of all other vascular plants and some primitive ferns, the initials are in a layer (i.e., more than one). A eusporangium is larger (hence contain more spores), and its wall is multi-layered. Although the wall may be stretched and damaged, resulting in only one cell-layer remaining.
A cluster of sporangia that have become fused in development is called a synangium. This structure is most prominent in ''Psilotum
'' and Marattiaceae
such as ''Christensenia
'' and ''Marattia
A columella (pl. columellae) is a sterile (non-reproductive) structure that extends into and supports the sporangium of some species. In fungi, the columella, which may be branched or unbranched, may be of fungal or host origin. ''Secotium
'' species have a simple, unbranched columella, while in ''Gymnoglossum
'' species, the columella is branched. In some ''Geastrum
'' species, the columella appears as an extension of the stalk into the spore mass (gleba
Category:Fungal morphology and anatomy