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Temporal range: CarboniferousPresent
Various gymnosperms.
Scientific classification Red Pencil Icon.png
Kingdom: Plantae
Clade: Tracheophytes
Clade: Spermatophytes
(unranked): Gymnosperms

Extant: Pinophyta (or Coniferophyta) – Conifers
Cycadophyta – Cycads
GnetophytaGnetum, Ephedra, Welwitschia


Extinct: Pteridospermatophyta - seed ferns

Encephalartos sclavoi cone, about 30 cm long Encephalartos sclavoi reproductive cone.jpg
Encephalartos sclavoi cone, about 30 cm long

The gymnosperms ( Loudspeaker.svg pronunciation   lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, Ginkgo , and gnetophytes, forming the clade Gymnospermae, the living members of which are also known as Acrogymnospermae. The term gymnosperm comes from the composite word in Greek : γυμνόσπερμος (γυμνός, gymnos, 'naked' and σπέρμα, sperma, 'seed'), literally meaning 'naked seeds'. The name is based on the unenclosed condition of their seeds (called ovules in their unfertilized state). The non-encased condition of their seeds contrasts with the seeds and ovules of flowering plants (angiosperms), which are enclosed within an ovary. Gymnosperm seeds develop either on the surface of scales or leaves, which are often modified to form cones, or solitary as in yew, Torreya , Ginkgo . [1] Gymnosperm lifecycles involve alternation of generations. They have a dominant diploid sporophyte phase and a reduced haploid gametophyte phase which is dependent on the sporophytic phase.

The gymnosperms and angiosperms together comprise the spermatophytes or seed plants. The gymnosperms are subdivided into five Divisions, four of which, the Cycadophyta, Ginkgophyta, Gnetophyta, and Pinophyta (also known as Coniferophyta) are still in existence while the Pteridospermatophyta are now extinct. [2]

By far the largest group of living gymnosperms are the conifers (pines, cypresses, and relatives), followed by cycads, gnetophytes ( Gnetum , Ephedra and Welwitschia ), and Ginkgo biloba (a single living species). About 65% of gymnosperms are dioecious, [3] but conifers are almost all monoecious. [4]

Some genera have mycorrhiza, fungal associations with roots (Pinus), while in some others (Cycas) small specialised roots called coralloid roots are associated with nitrogen-fixing cyanobacteria.


A formal classification of the living gymnosperms is the "Acrogymnospermae", which form a monophyletic group within the spermatophytes. [5] [6] The wider "Gymnospermae" group includes extinct gymnosperms and is thought to be paraphyletic. The fossil record of gymnosperms includes many distinctive taxa that do not belong to the four modern groups, including seed-bearing trees that have a somewhat fern-like vegetative morphology (the so-called "seed ferns" or pteridosperms). [7] When fossil gymnosperms such as these and the Bennettitales, glossopterids, and Caytonia are considered, it is clear that angiosperms are nested within a larger gymnospermae clade, although which group of gymnosperms is their closest relative remains unclear.

The extant gymnosperms include 12 main families and 83 genera which contain more than 1000 known species. [1] [6] [8]

Subclass Cycadidae

Subclass Ginkgoidae

Subclass Gnetidae

Subclass Pinidae

Extinct groupings

Diversity and origin

Over 1000 living species of gymnosperm exist. [1] It was previously widely accepted that the gymnosperms originated in the Late Carboniferous period, replacing the lycopsid rainforests of the tropical region, but more recent phylogenetic evidence indicates that they diverged from the ancestors of angiosperms during the Early Carboniferous. [9] [10] The radiation of gymnosperms during the late Carboniferous appears to have resulted from a whole genome duplication event around 319  million years ago. [11] Early characteristics of seed plants are evident in fossil progymnosperms of the late Devonian period around 383 million years ago. It has been suggested that during the mid-Mesozoic era, pollination of some extinct groups of gymnosperms was by extinct species of scorpionflies that had specialized proboscis for feeding on pollination drops. The scorpionflies likely engaged in pollination mutualisms with gymnosperms, long before the similar and independent coevolution of nectar-feeding insects on angiosperms. [12] [13] Evidence has also been found that mid-Mesozoic gymnosperms were pollinated by Kalligrammatid lacewings, a now-extinct family with members which (in an example of convergent evolution) resembled the modern butterflies that arose far later. [14]

Zamia integrifolia, a cycad native to Florida Zamia integrifolia02.jpg
Zamia integrifolia, a cycad native to Florida

Conifers are by far the most abundant extant group of gymnosperms with six to eight families, with a total of 65–70 genera and 600–630 species (696 accepted names). [15] Conifers are woody plants and most are evergreens. [16] The leaves of many conifers are long, thin and needle-like, other species, including most Cupressaceae and some Podocarpaceae, have flat, triangular scale-like leaves. Agathis in Araucariaceae and Nageia in Podocarpaceae have broad, flat strap-shaped leaves.

Cycads are the next most abundant group of gymnosperms, with two or three families, 11 genera, and approximately 338 species. A majority of cycads are native to tropical climates and are most abundantly found in regions near the equator. The other extant groups are the 95–100 species of Gnetales and one species of Ginkgo. [2]







Gymnosperms have major economic uses. Pine, fir, spruce, and cedar are all examples of conifers that are used for lumber, paper production, and resin. Some other common uses for gymnosperms are soap, varnish, nail polish, food, gum, and perfumes. [17]

Life cycle

Example of gymnosperm lifecycle Gymnosperm life cycle (en).png
Example of gymnosperm lifecycle

Gymnosperms, like all vascular plants, have a sporophyte-dominant life cycle, which means they spend most of their life cycle with diploid cells, while the gametophyte (gamete-bearing phase) is relatively short-lived. Like all seed plants, they are heterosporous, having two spore types, microspores (male) and megaspores (female) that are typically produced in pollen cones or ovulate cones, respectively. As with all heterosporous plants, the gametophytes develop within the spore wall. Pollen grains (microgametophytes) mature from microspores, and ultimately produce sperm cells. Megagametophytes develop from megaspores and are retained within the ovule. Gymnosperms produce multiple archegonia, which produce the female gamete. During pollination, pollen grains are physically transferred between plants from the pollen cone to the ovule. Pollen is usually moved by wind or insects. Whole grains enter each ovule through a microscopic gap in the ovule coat (integument) called the micropyle. The pollen grains mature further inside the ovule and produce sperm cells. Two main modes of fertilization are found in gymnosperms. Cycads and Ginkgo have flagellated motile sperm [18] that swim directly to the egg inside the ovule, whereas conifers and gnetophytes have sperm with no flagella that are moved along a pollen tube to the egg. After syngamy (joining of the sperm and egg cell), the zygote develops into an embryo (young sporophyte). More than one embryo is usually initiated in each gymnosperm seed. The mature seed comprises the embryo and the remains of the female gametophyte, which serves as a food supply, and the seed coat. [19]


The first published sequenced genome for any gymnosperm was the genome of Picea abies in 2013. [20]

Related Research Articles

Gametophyte 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.

Flowering plant Clade of seed plants that produce flowers

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 angeion and sperma ('seed'), and refers to those plants that produce their seeds enclosed within a fruit. They are the most diverse group of land plants with 64 orders, 416 families, approximately 13,000 known genera and 300,000 known species. Angiosperms were formerly called Magnoliophyta.

Fertilisation Union of gametes of opposite sexes during the process of sexual reproduction to form a zygote

Fertilisation or fertilization, 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. Processes such as insemination or pollination which happen before the fusion of gametes are also sometimes informally called fertilisation. The cycle of fertilisation and development of new individuals is called sexual reproduction. During double fertilisation in angiosperms the haploid male gamete combines with two haploid polar nuclei to form a triploid primary endosperm nucleus by the process of vegetative fertilisation.

Ginkgoales Order of plants

Ginkgoales are a gymnosperm order containing only one extant species: Ginkgo biloba, the ginkgo tree. It is monotypic, within the class Ginkgoopsida, which itself is monotypic within the division Ginkgophyta. The order includes five families, of which only Ginkgoaceae remains extant.

Conifer Group of cone-bearing seed plants

Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta, also known as Coniferophyta or Coniferae. The division contains a single extant class, Pinopsida. All extant conifers are perennial woody plants with secondary growth. The great majority are trees, though a few are shrubs. Examples include cedars, Douglas-firs, cypresses, firs, junipers, kauri, larches, pines, hemlocks, redwoods, spruces, and yews. As of 1998, the division Pinophyta was estimated to contain eight families, 68 genera, and 629 living species.

Pinales Order of seed plants, also known as conifers

The order Pinales in the division Pinophyta, class Pinopsida, comprises all the extant conifers. The distinguishing characteristic is the reproductive structure known as a cone produced by all Pinales. All of the extant conifers, such as cedar, celery-pine, cypress, fir, juniper, larch, pine, redwood, spruce, yew and Araucaria araucana are included here. Some fossil conifers, however, belong to other distinct orders within the division Pinophyta.

Archegonium Organ of the gametophyte of certain plants, producing and containing the ovum

An archegonium, from the ancient Greek ἀρχή ("beginning") and γόνος ("offspring"), is a multicellular structure or organ of the gametophyte phase of certain plants, producing and containing the ovum or female gamete. The corresponding male organ is called the antheridium. The archegonium has a long neck canal or venter and a swollen base. Archegonia are typically located on the surface of the plant thallus, although in the hornworts they are embedded.

Gnetophyta Division of plants containing three genera of gymnosperms

Gnetophyta is a division of plants, grouped within the gymnosperms, that consists of some 70 species across the three relict genera: Gnetum, Welwitschia, and Ephedra. Fossilized pollen attributed to a close relative of Ephedra has been dated as far back as the Early Cretaceous. Though diverse in the Early Cretaceous, only three families, each containing a single genus, are still alive today. The primary difference between gnetophytes and other gymnosperms is the presence of vessel elements, a system of conduits that transport water within the plant, similar to those found in flowering plants. Because of this, gnetophytes were once thought to be the closest gymnosperm relatives to flowering plants, but more recent molecular studies have brought this hypothesis into question.

Embryophyte Subclade of green plants, also known as land plants

The Embryophyta, or land plants, are the most familiar group of green plants that comprise vegetation on Earth. Embryophyta is a clade within the Phragmoplastophyta, a larger clade that also includes several groups of green algae including the Charophyceae and Coleochaetales. Within this larger clade the embryophytes are sister to the Zygnematophyceae/Mesotaeniaceae and consist of the bryophytes plus the polysporangiophytes. Living embryophytes therefore include hornworts, liverworts, mosses, lycophytes, ferns, gymnosperms and flowering plants.

Pollen tube A tubular structure to conduct male gametes of plants to the female gametes

A pollen tube is a tubular structure produced by the male gametophyte of seed plants when it germinates. Pollen tube elongation is an integral stage in the plant life cycle. The pollen tube acts as a conduit to transport the male gamete cells from the pollen grain—either from the stigma to the ovules at the base of the pistil or directly through ovule tissue in some gymnosperms. In maize, this single cell can grow longer than 12 inches (30 cm) to traverse the length of the pistil.

Sporophyte 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. It develops from the zygote produced when a haploid egg cell is fertilized by a haploid sperm and each sporophyte cell therefore has a double set of chromosomes, one set from each parent. All land plants, and most multicellular algae, have life cycles in which a multicellular diploid sporophyte phase alternates with a multicellular haploid gametophyte phase. In the seed plants, the largest groups of which are the gymnosperms and flowering plants (angiosperms), the sporophyte phase is more prominent than the gametophyte, and is the familiar green plant with its roots, stem, leaves and cones or flowers. In flowering plants the gametophytes are very reduced in size, and are represented by the germinated pollen and the embryo sac.

Ovule Female plant reproductive structure

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 female gametophyte in its center. The female gametophyte — specifically termed a megagametophyte— is also called the embryo sac in angiosperms. The megagametophyte produces an egg cell for the purpose of fertilization. The ovule is a small structure present in the ovary. It is attached to the placenta by a stalk called a funicle. The funicle provides nourishment to the ovule.

Endosperm Starchy tissue inside cereals and alike

The endosperm is a tissue produced inside the seeds of most of the flowering plants following double fertilization. It is triploid in most species, which may be auxin-driven. It surrounds the embryo and provides nutrition in the form of starch, though it can also contain oils and protein. This can make endosperm a source of nutrition in animal diet. For example, wheat endosperm is ground into flour for bread, while barley endosperm is the main source of sugars for beer production. Other examples of endosperm that forms the bulk of the edible portion are coconut "meat" and coconut "water", and corn. Some plants, such as orchids, lack endosperm in their seeds.

Plant reproductive morphology Study of the physical form and structure (the morphology) of those parts of plants directly or indirectly concerned with sexual reproduction

Plant reproductive morphology is the study of the physical form and structure of those parts of plants directly or indirectly concerned with sexual reproduction.

Bennettitales Extinct order of seed plants

Bennettitales is an extinct order of seed plants that first appeared in the Permian period and became extinct in most areas toward the end of the Cretaceous. Bennettitales are among the most common Mesozoic seed plants, and had morphologies including shrub and cycad-like forms. The foliage of bennettitaleans is superficially nearly indistinguishable from that of cycads, but they are distinguished from cycads by their more complex flower-like reproductive organs, at least some of which were likely pollinated by insects.

Double fertilization Complex fertilization mechanism of flowering plants

Double fertilization 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 megagametophyte.

Microspore 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. The resulting clonal plants are genetically identical to the parent plant and each other, unless mutations occur.

<i>Ginkgo</i> Genus of ancient seed plants with a single surviving species

Ginkgo is a genus of non-flowering seed plants. The scientific name is also used as the English name. The order to which it belongs, Ginkgoales, first appeared in the Permian, 270 million years ago, and is now the only living genus within the order. The rate of evolution within the genus has been slow, and almost all its species had become extinct by the end of the Pliocene. The sole surviving species, Ginkgo biloba is only found in the wild in China, but is cultivated around the world. The relationships between ginkgos and other groups of plants are not fully resolved.

Spermatophyte Clade of seed plants

A spermatophyte, also known as phanerogam or phaenogam, is any plant that produces seeds, hence the alternative name seed plant. Spermatophytes are a subset of the embryophytes or land plants.


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General bibliography