Strobilus
Encyclopedia
A strobilus is a structure present on many land plant
species consisting of sporangia
-bearing structures densely aggregated along a stem. Strobili are often called cones, but many botanists restrict the use of the term cone to the woody seed strobili of conifers. Strobili are characterized by a central axis (anatomically a stem
) surrounded by spirally arranged or decussate structures that may be modified leaves or modified stems.
Leaves that bear sporangia are called sporophylls, while sporangia bearing stems are called sporangiophores.
(Lycopodiopsida
and Isoetopsida
) produce strobili. In all cases, the lateral organs of the strobilus are microphyll
s, bearing sporangia
. In other lycophytes, ordinary foliage leaves can act as sporophyll
s, and there are no organized strobili.
s and pollen
in different structures. Strobili bearing microsporangia are called microsporangiate strobili or pollen cones, and those bearing ovules are megasporangiate strobili or seed cones (or ovulate cones).
are similar to those of cycads (although much smaller) and Ginkgoes in that they are composed of microsporophylls with microsporangia on the abaxial surface. Seed cones of many conifers are compound strobili. The central stem produces bracts and in the axil of each bract is a cone scale. Morphologically the cone scale is a reduced stem. Ovules are produced on the adaxial surface of the cone scales.
consists of three genera, Ephedra
, Gnetum
and Welwitschia
. All three are typically dioecious, although some Ephedra species exhibit monoecy. In contrast to the conifers, which have simple pollen strobili and compound seed strobili, gnetophytes have both compound pollen and seed strobili. The seed strobili of Ephedra and Gnetum are reduced, with Ephedra producing only two ovules per strobilus and Gnetum a single ovule.
of flowering plants is sometimes referred to as a bisexual strobilus. Stamen
s include microsporangia
within the anther, and ovule
s (contained in carpels) are megasporangia. Magnolia
has a particularly strobiloid flower with all parts arranged in a spiral, rather than as clear whorls.
A number of flowering plants have inflorescence
s that resemble strobili, such as catkin
s, but are actually more complex in structure than strobili.
is not unusual, since the form of a strobilus is one of the most compact that can be achieved in arranging lateral organs around a cylindric axis, and the consolidation of reproductive parts in a strobilus may optimize spore dispersal
and nutrient partitioning.
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...
species consisting of sporangia
Sporangium
A sporangium is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. All plants, fungi, and many other lineages form sporangia at some point in their life cycle...
-bearing structures densely aggregated along a stem. Strobili are often called cones, but many botanists restrict the use of the term cone to the woody seed strobili of conifers. Strobili are characterized by a central axis (anatomically a stem
Plant stem
A stem is one of two main structural axes of a vascular plant. The stem is normally divided into nodes and internodes, the nodes hold buds which grow into one or more leaves, inflorescence , conifer cones, roots, other stems etc. The internodes distance one node from another...
) surrounded by spirally arranged or decussate structures that may be modified leaves or modified stems.
Leaves that bear sporangia are called sporophylls, while sporangia bearing stems are called sporangiophores.
Lycophytes
Some members of both of the two modern classes of LycopodiophytaLycopodiophyta
The Division Lycopodiophyta is a tracheophyte subdivision of the Kingdom Plantae. It is the oldest extant vascular plant division at around 410 million years old, and includes some of the most "primitive" extant species...
(Lycopodiopsida
Lycopodiopsida
Lycopodiopsida is a class of plants often loosely grouped as the fern allies. Traditionally the group included not only the clubmosses and firmosses, but also the spikemosses and the quillworts...
and Isoetopsida
Isoetopsida
The Isoetopsida is a class of the Lycopodiophyta. All living plants belong to the genus Selaginella in the Selaginellales or to Isoetes in the order Isoetales. In the past, members of this group have sometimes been placed in the class Isoetopsida, sometimes in the Selaginellopsida or Lycopodiopsida...
) produce strobili. In all cases, the lateral organs of the strobilus are microphyll
Microphyll
The terminology of fossil plants is in places a little confusing. In the discipline's 200+ year history, certain concepts have become entrenched, even though improved understanding has threatened the foundations upon which they are based...
s, bearing sporangia
Sporangium
A sporangium is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. All plants, fungi, and many other lineages form sporangia at some point in their life cycle...
. In other lycophytes, ordinary foliage leaves can act as sporophyll
Sporophyll
A sporophyll is a leaf that bears sporangia. Both microphylls and megaphylls can be sporophylls. In heterosporous plants, sporophylls bear either megasporangia , or microsporangia...
s, and there are no organized strobili.
Sphenophytes
The single extant genus of Equisetophyta, Equisetum, produces strobili in which the lateral organs are sporangiophores. Developmental evidence and comparison with fossil members of the group show that the sporangiophores are reduced stems, rather than leaves. Sporangia are terminal.Seed plants
With the exception of flowering plants, seed plants produce ovuleOvule
Ovule means "small egg". 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 , and the megaspore-derived female gametophyte in its center...
s and pollen
Pollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...
in different structures. Strobili bearing microsporangia are called microsporangiate strobili or pollen cones, and those bearing ovules are megasporangiate strobili or seed cones (or ovulate cones).
Cycads
Cycadophyta are typically dioecious (seed strobili and pollen strobili are produced on separate plants). The lateral organs of seed strobili are megasporophylls (modified leaves) that bear two to several marginal ovules. Pollen strobili consist of microsporophylls, each of which may have dozens or hundreds of abaxial microsporangia.Ginkgoes
The single living member of the Ginkgophyta, Ginkgo biloba produces pollen strobili, but the ovules are borne in pairs at the end of a stem, not in a strobilus.Conifers
Pollen strobili of PinophytaPinophyta
The conifers, division Pinophyta, also known as division Coniferophyta or Coniferae, are one of 13 or 14 division level taxa within the Kingdom Plantae. Pinophytes are gymnosperms. They are cone-bearing seed plants with vascular tissue; all extant conifers are woody plants, the great majority being...
are similar to those of cycads (although much smaller) and Ginkgoes in that they are composed of microsporophylls with microsporangia on the abaxial surface. Seed cones of many conifers are compound strobili. The central stem produces bracts and in the axil of each bract is a cone scale. Morphologically the cone scale is a reduced stem. Ovules are produced on the adaxial surface of the cone scales.
Gnetophytes
GnetophytaGnetophyta
The plant division Gnetophyta or gnetophytes consists of three genera of woody plants grouped in the gymnosperms. The living Gnetophyta consists of around 70 species across the three genera Gnetum , Welwitschia , and Ephedra .The gnetophytes differ from other gymnosperms The plant division...
consists of three genera, Ephedra
Ephedra (genus)
Ephedra is a genus of gymnosperm shrubs, the only genus in its family, Ephedraceae, and order, Ephedrales. Ephedra grows in dry climates over wide areas of the northern hemisphere, including southwestern North America, Europe, north Africa, and southwest and central Asia, and, in the southern...
, Gnetum
Gnetum
Gnetum is a genus of about 30-35 species of gymnosperms, the sole genus in the family Gnetaceae and order Gnetales. They are tropical evergreen trees, shrubs and lianas. Unlike other gymnosperms they possess vessel elements in the xylem...
and Welwitschia
Welwitschia
Welwitschia is a monotypic genus of gymnosperm plant, composed solely of the very distinct Welwitschia mirabilis. The plant is commonly simply known as Welwitschia in English. It is known locally as !kharos or khurub , tweeblaarkanniedood , nyanka , or onyanga , among others...
. All three are typically dioecious, although some Ephedra species exhibit monoecy. In contrast to the conifers, which have simple pollen strobili and compound seed strobili, gnetophytes have both compound pollen and seed strobili. The seed strobili of Ephedra and Gnetum are reduced, with Ephedra producing only two ovules per strobilus and Gnetum a single ovule.
Flowering plants
The flowerFlower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants . The biological function of a flower is to effect reproduction, usually by providing a mechanism for the union of sperm with eggs...
of flowering plants is sometimes referred to as a bisexual strobilus. Stamen
Stamen
The stamen is the pollen producing reproductive organ of a flower...
s include microsporangia
Microsporangia
Microsporangium is a sporangium that produces spores that give rise to male gametophytes. Microsporangia are notable in spikemosses, and a minority of ferns. In Gymnosperms and Angiosperms , the microsporangium produce the microsporocyte, also known as the microspore mother cell, which then creates...
within the anther, and ovule
Ovule
Ovule means "small egg". 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 , and the megaspore-derived female gametophyte in its center...
s (contained in carpels) are megasporangia. Magnolia
Magnolia
Magnolia is a large genus of about 210 flowering plant species in the subfamily Magnolioideae of the family Magnoliaceae. It is named after French botanist Pierre Magnol....
has a particularly strobiloid flower with all parts arranged in a spiral, rather than as clear whorls.
A number of flowering plants have inflorescence
Inflorescence
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Strictly, it is the part of the shoot of seed plants where flowers are formed and which is accordingly modified...
s that resemble strobili, such as catkin
Catkin
A catkin or ament is a slim, cylindrical flower cluster, with inconspicuous or no petals, usually wind-pollinated but sometimes insect pollinated . They contain many, usually unisexual flowers, arranged closely along a central stem which is often drooping...
s, but are actually more complex in structure than strobili.
Evolution of strobili
It is likely that strobili evolved independently in most if not all these groups. This evolutionary convergenceConvergent evolution
Convergent evolution describes the acquisition of the same biological trait in unrelated lineages.The wing is a classic example of convergent evolution in action. Although their last common ancestor did not have wings, both birds and bats do, and are capable of powered flight. The wings are...
is not unusual, since the form of a strobilus is one of the most compact that can be achieved in arranging lateral organs around a cylindric axis, and the consolidation of reproductive parts in a strobilus may optimize spore dispersal
Biological dispersal
Biological dispersal refers to species movement away from an existing population or away from the parent organism. Through simply moving from one habitat patch to another, the dispersal of an individual has consequences not only for individual fitness, but also for population dynamics, population...
and nutrient partitioning.