Gymnosperms are seed plants that do not produce flowers. The term gymnosperm means "naked seed." However, usually when the seeds of gymnosperms are immature they are enclosed within and protected by modified leaves or a cone. In flowering plants (or angiosperms, which means "vessel seed") the ovary wall or fruit encloses the seeds, whereas in gymnosperms there is no equivalent structure; hence, the interpretation of the seeds as "naked" or not enclosed.
There are four groups of gymnosperms living today—Coniferophyta, Cycadophyta, Ginkgophyta, and Gnetophyta—but many additional groups are known from the fossil record. Seed plants evolved more than 350 million years ago and the first seed plants were gymnosperms. The relationship of the flowering plants to the gymnospermous seed plants remains a hotly contested issue within the scientific community.
Although each group of gymnosperms has its own specific characteristics, some features are shared throughout. For example, all gymnosperms produce at least some secondary growth, whereas secondary growth is lacking from most spore-bearing (nonseed) plants. Secondary growth is plant growth that does not occur directly from the tips of the plant; it is growth that occurs horizontally (or radially) rather than vertically. The most abundant product of secondary growth is the secondary xylem (xylem is the water-conducting tissue of plants), or wood.
Aspects of reproduction are also shared by all the gymnosperms. The ovule (the technical term for the seed prior to fertilization) consists of female nutritive tissue plus the female gamete, or egg, both enclosed by a
|DISTINGUISHING CHARACTERS OF GYMNOSPERMS|
|Extant members||50-60 genera, about 550 species||11 genera, about 160 species||1 species (Ginkgo biloba)||3 genera: Ephedra (35 species), Gnetum (30 species), Welwitschia (1 species)|
|Distribution||Worldwide, especially temperate regions||Tropics||Native to a small area of China||Isolated areas of temperate and tropical regions|
|Habit||Trees with branched woody trunk||Trees with unbranched fleshy trunk||Trees with branched woody trunk||Shrubs, vines, small trees, or weird tubers|
|Leaves||Simple, needlelike or broad||Large, compound (each leaf composed of many leaflets)||Simple, fan-shaped||Simple, needlelike or broad|
|Reproductive structure||Simple (unbranched) male, compound (branched) female cones||Simple cones (unbranched), male or female||Simple cones (unbranched), male or female||Compound (branched) male or female cones|
|Sex||Separate male and female cones on each plant (monoecious) or separate male and female plants (dioecious)||Individual plant male or female (dioecious)||Individual plant male or female (dioecious)||Individual plant male or female (dioecious)|
|Seeds||Small, without differentiated layers||Large, with outer fleshy layer and middle stony layer||Large, with outer fleshy layer and middle stony layer||Small, without differentiated layers|
|Sperm||No flagella; pollen tube delivers to egg||Swims using many flagella||Swims using many flagella||No flagella; pollen tube delivers to egg|
layer of protective tissue (the integument). The male gamete is carried within the pollen. In most gymnosperms pollen is produced in great amounts, and the pollen grains are dispersed by the wind. Usually a small amount of sticky liquid (the pollination drop) is exuded at the tip of the ovule. Pollen grains become stuck in the pollination drop and, as the drop dries, the pollen is pulled into the ovule. Depending on the type of gymnosperm, the male ga-mete is released from the pollen and swims to the egg, or the male gamete is transported within a tube that grows to the egg. The fertilized egg then develops into the embryo of the seed. Ultimately, when the seed germinates, the embryo grows to produce the young seedling using the female nutritive tissue as a source of energy. An unusual attribute of the gymnosperms, except for the gnetophytes, is the long length of time—a year or more—that passes between the production of the egg and the sperm and the actual occurrence of fertilization.
The Coniferophyta, or conifers, are the most abundant group of living gymnosperms and the first of the living gymnosperm groups to appear in the fossil record. They have been important components of Earth's vegetation for almost three hundred million years. The oldest (bristlecone pine), the tallest (coast redwood), and the biggest (giant sequoia) organisms on the planet today are conifers. Most conifers are large trees that make abundant wood, have small evergreen leaves, and produce their seeds within woody cones.
Although most conifers fit that standard description, there are numerous exceptions. First, not all conifers are evergreen. A few types, such as the larch, are deciduous, losing all their leaves each fall and growing new leaves in the spring. Even the leaves of the evergreen type are not immortal; it is just that the senescence (aging and death) of leaves occurs individually rather than all at once. Second, not all conifers have small or needlelike leaves. Some conifers, native to the Southern Hemisphere, have broad leaves. Third, although all conifers are woody plants, not all conifers are large trees. A number are shrubs or small trees, and one is even a parasite on the roots of other coniferous trees. Finally, not all conifers have cones as people usually think of them. Some conifers, such as the yew, have solitary seeds covered with a fleshy, colorful tissue.
The female cones produced by most conifers are complex structures that are made of repeating units, each consisting of ovules on a woody platform (ovuliferous scale) beneath which is a bract (e.g., the "mouse tail" that peeks out of Douglas-fir cones). Cones of the other gymnosperms lack the equivalent of the ovuliferous scale. Conifers make rather small ovules and lack swimming sperm.
The Cycadophyta, or cycads, are restricted to tropical latitudes and were more abundant in the geologic past. These plants have unbranched, fleshy stems. The trunks of some species can grow fairly tall (15 to 18 meters), but all cycads lack extensive wood development. Because so much water is present in the stems, the plants are very vulnerable to damage from freezing (think of a soda can that explodes after you forget you have placed it in the freezer). Thus the cycads are restricted to parts of the world where freezing temperatures are rare or absent. A few types are found naturally in sub-tropical areas where mild freezing temperatures occasionally occur; these types of cycads have short, squat, subterranean stems. Some additional cycad species are hardy as ornamentals in similar subtropical to warm temperate areas. Excellent outdoor collections of cycads can be seen in the United States at Fairchild Botanical Garden in Miami, Florida, or at Huntington Botanical Garden east of Los Angeles, California.
The leaves of cycads occur as a crown at the top of the stem. Most often the leaves are very large (up to 1.5 meters in length), leathery, and compound; that is, the blade of the leaf is made of many separate leaflets. When the leaves fall off, usually the base of the leaf remains attached to the stem. Thus, the trunk is protected from herbivory and to some extent from freezing by the remnant bases.
Individual cycad plants are either male or female. At the tip of the trunk within the crown of leaves a cone develops that will either contain pollen or ovules, depending on the sex of the plant. Cycads make very large ovules and swimming sperm.
Only one species of the Ginkgophyta group remains living today: the maidenhair tree, Ginkgo biloba. Twenty to thirty million years ago the ancestor of the modern species was found throughout the Northern Hemisphere. However, climatic changes have led to the plants gradually becoming reduced to a smaller and smaller territory. Since the last ice age, wild populations have been restricted to a small area in China. It is debated whether or not any populations are truly wild. The tree still exists today because it was a sacred plant maintained in monasteries. Now the ginkgo tree is commonly used as a street tree and so has regained, in some sense, much of its earlier territory. Individual plants can live very long lives (more than one thousand years), and the species is unusually resistant to pollution and disease. The Ginkgo plant is a fairly tall (25 meters or more), much-branched tree with abundant wood, bearing fan-shaped leaves.
Individual ginkgo trees are either male or female. The sex of the plant does not become apparent until the tree is fifteen to twenty years old. Male plants are preferred as ornamentals. The ovule produced by the female is large with a stony interior and a fleshy outer covering that produces a smell usually equated with rotting butter. Ginkgo also produces swimming sperm.
The Gnetophyta are a small, odd group of living gymnosperms. Three types are known, each being somewhat different from the other gnetophytes and unusual among the seed plants in its own way. Species of the genus Ephedra are shrubby plants of arid temperate regions. They have tiny leaves and the stems are often green and photosynthetic. Species of the genus Gnetum are tropical plants that are small trees or, more usually, vines. The gnetums are unusual among the gymnosperms in that they produce broad leaves with netted venation similar to that found in flowering plants. The third group is represented by a single species, Welwitschia mirabilis. Welwitschia has been described as the "weirdest plant on Earth," a plant that has "lost its head" or a giant seedling that can reproduce. Welwitschia is native to the Namib Desert of southwest Africa and consists of a large tuber or taproot and two long, strap-shaped leaves that grow from their bases and are retained for the life of the plant.
Most gnetophytes produce separate female and male cones born on separate (male or female) plants. Each type of gnetophyte has a slightly different reproductive structure but all are variations on one theme: pairs of papery bracts (modified leaves) surround the ovule or pollen organs. Some botanists compare these bracts to the petals and sepals of angiosperm flowers, whereas others equate them to components of the cone found in conifers. Other aspects of the gnetophyte morphology are also ambiguous and, depending on their interpretation, suggests an evolutionary link to either the conifers or the angiosperms. For example, vessels, specialized water-conducting cells, occur in the wood of gnetophytes. These cells are dead at maturity, empty of any contents. Vessels lack any end walls; they are open-ended tubes that line up end to end and act as a pipe to transport water without any obstructions. Most flowering plants have vessels but most other types of plants lack this cell type. Some plant scientists see the gnetophyte vessels as evidence that gnetophytes and angiosperms share a common ancestor. However, because of differences in vessel cell anatomy and development between gnetophytes and flowering plants, other botanists consider the gnetophyte vessel to be an independent evolution of a water transport cell that lost an end wall. Molecular comparisons of the seed plants using different types of deoxyribonucleic acid (DNA) data have not settled this debate: gnetophytes remain a problematic group in terms of evolutionary placement and morphological interpretation.
Conifers are the gymnospermous group with the most profound ecological role in Earth's vegetation. Conifers dominate some vegetation types, such as the taiga of high northern latitudes or boreal forests of lower latitudes. Some temperate forests, for example in Argentina, Australia, or northwestern North America, are also composed almost exclusively of coniferous trees.
Conifers can also be important as successional species or as the climax vegetation of odd environments in other temperate or tropical areas. For example, in environments prone to fire, with a long enough growing season and enough precipitation to support the growth of trees, conifers are often present. Conifer bark is thicker than most flowering plant tree bark and so coniferous trees are better able to survive ground fires. Some conifer cones open to release their seeds only after a fire.
Conifers are also considered the most important gymnospermous group from an economic perspective. Coniferous trees are a very important source of timber for lumber and paper. They are harvested in North America, parts of Europe and Asia, and in Australia. In addition to timber, conifers provide Christmas trees, ornamental trees and shrubs, turpentine, and resin. Pine nuts (or pignoli), the seeds of some pine trees, are used as food. An important cancer-fighting drug, taxol, has been derived from the bark and leaves of the Pacific Coast yew (Taxus ). Other gymnosperms also are the source of drugs and herbal medications. The powerful stimulant ephedrine derived from the gnetophyte Ephedra is often used in cold and allergy medications, and compounds shown to improve the mental capacities of the elderly have been discovered in Ginkgo. Ginkgo seeds are also quite nutritious and are used as food in Asia. Ginkgo and cycads are also important as ornamentals.
see also Coniferous Forests; Conifers; Evolution of Plants; Ginkgo; Record-Holding Plants; Trees; Wood Products.
Linda A. Raubeson
Burns, Russell M., and Barbara H. Honkala. Silvics of North America, Vol. 1: Conifers. Washington, DC: USDA Forest Service, Agriculture Handbook 654, 1990.
Homan, Dennis. Biology of Seed Plants: A Laboratory Manual. Dubuque, IA:Kendall/Hunt Publishing Co., 1997.
Krussmann, Gerd. Manual of Cultivated Conifers. Portland, OR: Timber Press, 1985.
Nimsch, Hubertus. A Reference Guide to the Gymnosperms of the World. Champaign, IL: Balogh Scientific Books, 1995.
Norstog, Kurt J., and Trevor J. Nicholls. The Biology of the Cycads. Ithaca, NY: Cornell University Press, 1998.
Pielou, E. C. The World of Northern Evergreens. Ithaca, NY: Cornell University Press,1988.
Gymnosperms are a group of plants that share one common characteristic: they bear seeds, but their seeds do not develop within an ovary. For this reason, gymnosperms were long thought to be an evolutionary precursor to the angiosperms, which are seed plants that enclose their seeds in an ovary and that are vastly more diverse than gymnosperms. Studies of their deoxyribonucleic acid (DNA) has shown that the gymnosperms consist of four major, related groups: conifers, cycads, ginkgo, and gnetophytes.
With approximately 588 living species, this is the most diverse and by far the most ecologically and economically important gymnosperm group. Conifers grow in all climate zones and on all continents except Antarctica. They all bear their seeds within a cone or a structure superficially resembling a berry (true berries only exist among angiosperms). Most conifers are trees. Conifers appeared in the fossil record about 290 million years ago and have been an ecologically important, widespread group ever since then.
The 220 species of cycads are widely distributed through the tropical and subtropical regions. Most of them superficially resemble ferns, having a cluster of long pinnate (rarely bipinnate) fronds growing from a central stalk, but they differ in developing distinctive male and female cones. Cycads are woody, long-lived, unisexual plants. All species have coralloid roots, which support symbiotic cyanobacteria that can fix atmospheric nitrogen. The cycads and ginkgo are unique among seed plants in having motile sperm; this is often taken as evidence of their evolutionary primitiveness. Cycads appeared in the fossil record about 230 million years ago and attained their greatest ecological importance during the Jurassic period, about 193 million to 136 million years ago, when they formed extensive forests.
There is one surviving species of ginkgo. It is a tree, sometimes attaining large size, native to China but widely planted around the world. Ginkgo is often referred to as a "living fossil" because nearly identical plants are known from fossils nearly 200 million years old. The fossil record shows that they were formerly a widespread, abundant, and diverse group.
The gnetophytes are one of the most peculiar plant groups. They include three highly distinct groups totaling 68 species. One group, the genus Ephedra, is composed of shrubs native to deserts and semiarid areas. The second group, the genus Gnetum, is composed of climbing vines (and one tree species) native to tropical rainforests. The third group contains a single species, Welwitschia mirabilis. It lives in the desert of Southwest Africa, produces two leaves that grow throughout the life of plant, and lives an estimated two thousand years. Although the fossil record is virtually nonexistent, studies suggest that the Gnetales are a relatively young group that evolved from the angiosperms and thus are unrelated to the other gymnosperms.
see also Conifers; Cyanobacteria; Nitrogen Fixation; Plant
Christopher J. Earle
Earle, Christopher J. The Gymnosperm Database. 2001. <http://www.conifers.org>.
Farabee, M. J. On-Line Biology Book: Biological Diversity: Seed Plants. <http://gened.emc.maricopa.edu/bio/bio181/BIOBK/BioBookDiversity_6.html>.
Norstog, Knut J., and Trevor J. Nicholls. The Biology of the Cycads. Ithaca, NY: Cornell University Press, 1997.
Gymnosperms are one of the two major groups of plants that produce seeds; the other is the angio-sperms. Gymnosperm literally means “naked seed,” which refers to the development of seeds exposed on a flat structure, that is, not within an ovary as in the angiosperms.
Gymnosperms became common about 290 million years ago and although many of the earlier types are now extinct, four kinds remain alive: the conifers, cycads, gnetophytes, and ginkgo, the maidenhair tree. Conifers are the most familiar, widespread, and abundant of the gymnosperms. Most conifers have needle-or scale-like leaves that persist for more than one year, that is, are evergreen. The conifers include species of pine, spruce, fir, hemlock, cedar, juniper, and redwood. The latter is the largest plant, exceeding 328 ft (100 m) in height. Conifers dominate boreal forests of high latitudes and mature forests of high altitudes, and are extremely valuable commercially. Their wood is used to make houses, furniture, and paper products. Resin, an organic secretion of conifers, has various uses ranging from turpentine for thinning paint, to resin blocks for making violin bows sticky. Some pines produce edible nuts and juniper berries are used to give gin its distinctive taste.
The other kinds of living gymnosperms are much less abundant, more geographically restricted, and less valuable than conifers. These gymnosperms are highly diverse, however, and sometimes quite strange. The cycads, or Sago palms, look like palms but are not (palms are flowering plants). Cycads are unusual among plants in that each individual is either a male or female, as in most animals. Most seed-producing plants are bisexual. The gnetophytes as a group are the oddest of the gymnosperms, and the oddest of these is Welwitschia. This species lives in sandy deserts of southwestern Africa. Welwitschia has a saddlelike, central core that produces only two leaves during the life of the plant. These grow continuously, frequently splitting along their length to give a mop-headed appearance.
Lastly, the ginkgo or maidenhair tree (Ginkgo biloba ) is the only living species of a group that was most common about 170 million years ago. The ginkgo has distinctive, fan-shaped leaves that are sometimes cleft in the middle and, unlike the leaves of most gymnosperms, fall each autumn. Ginkgo trees also are either male or female. Because of its unusual and attractive leaves, this species is widely planted as an ornamental. Curiously, the ginkgo appears to be extinct in the wild, only having survived through cultivation by Buddhist monks in China.
Gymnosperms are one of the two major groups of plants that produce seeds ; the other is the angiosperms. Gymnosperm literally means "naked seed," which refers to the development of seeds exposed on a flat structure, that is, not within an ovary as in the angiosperms.
Gymnosperms became common about 290 million years ago and although many of the earlier types are now extinct, four kinds remain alive: the conifers, cycads , gnetophytes, and ginkgo , the maidenhair tree . Conifers are the most familiar, widespread, and abundant of the gymnosperms. Most conifers have needle- or scale-like leaves that persist for more than one year, that is, are evergreen. The conifers include species of pine, spruce , fir, hemlock, cedar, juniper , and redwood. The latter is the largest plant , exceeding 328 ft (100 m) in height. Conifers dominate boreal forests of high latitudes and mature forests of high altitudes, and are extremely valuable commercially. Their wood is used to make houses, furniture, and paper products. Resin, an organic secretion of conifers, has various uses ranging from turpentine for thinning paint, to resin blocks for making violin bows sticky. Some pines produce edible nuts and juniper berries are used to give gin its distinctive taste.
The other kinds of living gymnosperms are much less abundant, more geographically restricted, and less valuable than conifers. These gymnosperms are highly diverse, however, and sometimes quite strange. The cycads, or Sago palms , look like palms but are not (palms are flowering plants). Cycads are unusual among plants in that each individual is either a male or female, as in most animals. Most seed-producing plants are bisexual. The gnetophytes as a group are the oddest of the gymnosperms, and the oddest of these is Welwitschia. This species lives in sandy deserts of southwestern Africa . Welwitschia has a saddle-like, central core that produces only two leaves during the life of the plant. These grow continuously, frequently splitting along their length to give a mop-headed appearance.
Lastly, the ginkgo or maidenhair tree (Ginkgo biloba) is the only living species of a group that was most common about 170 millon years ago. The ginkgo has distinctive, fan-shaped leaves that are sometimes cleft in the middle and, unlike the leaves of most gymnosperms, fall each autumn. Ginkgo trees also are either male or female. Because of its unusual and attractive leaves, this species is widely planted as an ornamental. Curiously, the ginkgo appears to be extinct in the wild, only having survived through cultivation by Buddhist monks in China.