Sporophyte

A sporophyte (/spɔːroʊˌfaɪt/) 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, (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.

The sporophyte produces spores (hence the name) by meiosis, a process also known as "reduction division" that reduces the number of chromosomes in each spore mother cell by half. The resulting meiospores develop into a gametophyte. Both the spores and the resulting gametophyte are haploid, meaning they only have one set of chromosomes. The mature gametophyte produces male or female gametes (or both) by mitosis. The fusion of male and female gametes produces a diploid zygote which develops into a new sporophyte. This cycle is known as alternation of generations or alternation of phases.

Acer palmatum BotGartenMuenster Faecherahorn 6691
In flowering plants, the sporophyte comprises the whole multicellular body except the pollen and embryo sac

Bryophytes (mosses, liverworts and hornworts) have a dominant gametophyte phase on which the adult sporophyte is dependent for nutrition. The embryo sporophyte develops by cell division of the zygote within the female sex organ or archegonium, and in its early development is therefore nurtured by the gametophyte.[1] Because this embryo-nurturing feature of the life cycle is common to all land plants they are known collectively as the embryophytes.

Physcomitrella Sporophyt
Cleistocarpous sporophyte of the moss Physcomitrella patens

Most algae have dominant gametophyte generations, but in some species the gametophytes and sporophytes are morphologically similar (isomorphic). An independent sporophyte is the dominant form in all clubmosses, horsetails, ferns, gymnosperms, and angiosperms that have survived to the present day. Early land plants had sporophytes that produced identical spores (isosporous or homosporous) but the ancestors of the gymnosperms evolved complex heterosporous life cycles in which the spores producing male and female gametophytes were of different sizes, the female megaspores tending to be larger, and fewer in number, than the male microspores.

During the Devonian period several plant groups independently evolved heterospory and subsequently the habit of endospory, in which the gametophytes develop in miniaturized form inside the spore wall. By contrast in exosporous plants, including modern ferns, the gametophytes break the spore wall open on germination and develop outside it. The megagametophytes of endosporic plants such as the seed ferns developed within the sporangia of the parent sporophyte, producing a miniature multicellular female gametophyte complete with female sex organs, or archegonia. The oocytes were fertilized in the archegonia by free-swimming flagellate sperm produced by windborne miniaturized male gametophytes in the form of pre-pollen. The resulting zygote developed into the next sporophyte generation while still retained within the pre-ovule, the single large female meiospore or megaspore contained in the modified sporangium or nucellus of the parent sporophyte. The evolution of heterospory and endospory were among the earliest steps in the evolution of seeds of the kind produced by gymnosperms and angiosperms today.

Mech plonnik mlode sporofity
Young sporophytes of the common moss Tortula muralis. In mosses, the gametophyte is the dominant generation, while the sporophytes consist of sporangium-bearing stalks growing from the tips of the gametophytes
Macro Photography of Moss Sporophytes
Sporophytes of moss during spring

References

  1. ^ Ralf Reski(1998): Development, genetics and molecular biology of mosses. In: Botanica Acta. Bd. 111, S. 1-15.
  • P. Kenrick & P.R. Crane (1997) The origin and early evolution of plants on land. Nature 389, 33-39.
  • T.N. Taylor, H. Kerp and H. Hass (2005) Life history biology of early land plants: Deciphering the gametophyte phase. Proceedings of the National Academy of Sciences 102, 5892-5897.
  • P.R. Bell & A.R. Helmsley (2000) Green plants. Their Origin and Diversity. Cambridge University Press ISBN 0-521-64673-1
Alternation of generations

Alternation of generations (also known as metagenesis) is the type of life cycle that occurs in those plants and algae in the Archaeplastida and the Heterokontophyta that have distinct sexual haploid and asexual diploid stages. In these groups, a multicellular gametophyte, which is haploid with n chromosomes, alternates with a multicellular sporophyte, which is diploid with 2n chromosomes, made up of n pairs. A mature sporophyte produces spores by meiosis, a process which reduces the number of chromosomes to half, from 2n to n.

The haploid spores germinate and grow into a haploid gametophyte. At maturity, the gametophyte produces gametes by mitosis, which does not alter the number of chromosomes. Two gametes (originating from different organisms of the same species or from the same organism) fuse to produce a zygote, which develops into a diploid sporophyte. This cycle, from gametophyte to gametophyte (or equally from sporophyte to sporophyte), is the way in which all land plants and many algae undergo sexual reproduction.

The relationship between the sporophyte and gametophyte varies among different groups of plants. In those algae which have alternation of generations, the sporophyte and gametophyte are separate independent organisms, which may or may not have a similar appearance. In liverworts, mosses and hornworts, the sporophyte is less well developed than the gametophyte and is largely dependent on it. Although moss and hornwort sporophytes can photosynthesise, they require additional photosynthate from the gametophyte to sustain growth and spore development and depend on it for supply of water, mineral nutrients and nitrogen. By contrast, in all modern vascular plants the gametophyte is less well developed than the sporophyte, although their Devonian ancestors had gametophytes and sporophytes of approximately equivalent complexity. In ferns the gametophyte is a small flattened autotrophic prothallus on which the young sporophyte is briefly dependent for its nutrition. In flowering plants, the reduction of the gametophyte is much more extreme; it consists of just a few cells which grow entirely inside the sporophyte.

Animals develop differently. They directly produce haploid gametes. No haploid spores capable of dividing are produced, so they do not have a haploid gametophyte alternating with a diploid sporophyte. (Some insects have a sex-determining system whereby haploid males are produced from unfertilized eggs; however the females are diploid.)

Life cycles of plants and algae with alternating haploid and diploid multicellular stages are referred to as diplohaplontic (the equivalent terms haplodiplontic, diplobiontic or dibiontic are also in use). Life cycles, such as those of animals, in which there is only a diploid multicellular stage are referred to as diplontic. Life cycles in which there is only a haploid multicellular stage are referred to as haplontic.

Bryophyte

Bryophytes are an informal group consisting of three divisions of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. They 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 (gametangia and sporangia), but they do not produce flowers or seeds. They reproduce via spores. Bryophytes are usually considered to be a paraphyletic group and not a monophyletic group, although some studies have produced contrary results. Regardless of their status, the name is convenient and remains in use as an informal collective term. The term "bryophyte" comes from Greek βρύον, bryon "tree-moss, oyster-green" and φυτόν, phyton "plant".

The defining features of bryophytes are:

Their life cycles are dominated by the gametophyte stage

Their sporophytes are unbranched

They do not have a true vascular tissue containing lignin (although some have specialized tissues for the transport of water)

Cladophora

Cladophora is a genus of reticulated filamentous Ulvophyceae (green algae). The genus Cladophora contains many species that are very hard to tell apart and classify, mainly because of the great variation in their appearances, which is affected by habitat, age and environmental conditions. Unlike Spirogyra the filaments of Cladophora branch and do not undergo conjugation. There are two multicellular stages in its life cycle - a haploid gametophyte and a diploid sporophyte - which look highly similar. The only way to tell the two stages apart is to either count their chromosomes, or examine their offspring. The haploid gametophyte produces haploid gametes by mitosis and the diploid sporophyte produces haploid spores by meiosis. The only visible difference between the gametes and spores of Cladophora is that the gametes have two flagella and the spores have four. The Cladophora species can be a major nuisance causing major alteration to benthic conditions linked particularly with increased phosphorus loading.

Dioecy

Dioecy (Greek: διοικία "two households"; adjective form: dioecious) is a characteristic of a species, meaning that it has distinct male and female individual organisms. Dioecious reproduction is biparental reproduction. Dioecy is one method that excludes self-fertilization and promotes allogamy (outcrossing), and thus tends to reduce the expression of recessive deleterious mutations present in a population. Flowering plants have several other methods of excluding self-fertilization, called self-incompatibility.

Embryophyte

The Embryophyta, or land plants, are the most familiar group of green plants that form vegetation on earth. Embryophyta is a clade within the Phragmoplastophyta, a larger clade that also includes several green algae groups (including the Charophyceae and Coleochaetales), and within this large clade the embryophytes are sister to the Zygnematophyceae/Mesotaeniaceae and consist of the bryophytes plus the polysporangiophytes. Living embryophytes therefore include hornworts, liverworts, mosses, ferns, lycophytes, gymnosperms and flowering plants. The Embryophyta are informally called land plants because they live primarily in terrestrial habitats, while the related green algae are primarily aquatic. Embryophytes are complex multicellular eukaryotes with specialized reproductive organs. The name derives from their innovative characteristic of nurturing the young embryo sporophyte during the early stages of its multicellular development within the tissues of the parent gametophyte. With very few exceptions, embryophytes obtain their energy by photosynthesis, that is by using the energy of sunlight to synthesize their food from carbon dioxide and water.

Fern

A fern is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissues that conduct water and nutrients and in having life cycles in which the sporophyte is the dominant phase. Ferns have complex leaves called megaphylls, that are more complex than the microphylls of clubmosses. Most ferns are leptosporangiate ferns, sometimes referred to as true ferns. They produce coiled fiddleheads that uncoil and expand into fronds. The group includes about 10,560 known extant species.Ferns are defined here in the broad sense, being all of the Polypodiopsida, comprising both the leptosporangiate (Polypodiidae) and eusporangiate ferns, the latter itself comprising ferns other than those denominated true ferns, including horsetails or scouring rushes, whisk ferns, marattioid ferns, and ophioglossoid ferns.

Ferns first appear in the fossil record about 360 million years ago in the late Devonian period, but many of the current families and species did not appear until roughly 145 million years ago in the early Cretaceous, after flowering plants came to dominate many environments. The fern Osmunda claytoniana is a paramount example of evolutionary stasis; paleontological evidence indicates it has remained unchanged, even at the level of fossilized nuclei and chromosomes, for at least 180 million years.Ferns are not of major economic importance, but some are used for food, medicine, as biofertilizer, as ornamental plants and for remediating contaminated soil. They have been the subject of research for their ability to remove some chemical pollutants from the atmosphere. Some fern species, such as bracken (Pteridium aquilinum) and water fern (Azolla filiculoides) are significant weeds world wide. Some fern genera, such as Azolla can fix nitrogen and make a significant input to the nitrogen nutrition of rice paddies. They also play certain roles in mythology and art.

Gametophyte

A gametophyte () is one of the two alternating phases in the life cycle 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.

Hornwort

Hornworts are a group of non-vascular plants constituting the division Anthocerotophyta. The common name refers to the elongated horn-like structure, which is the sporophyte. As in mosses and liverworts, the flattened, green plant body of a hornwort is the gametophyte plant.

Hornworts may be found worldwide, though they tend to grow only in places that are damp or humid. Some species grow in large numbers as tiny weeds in the soil of gardens and cultivated fields. Large tropical and sub-tropical species of Dendroceros may be found growing on the bark of trees.

The total number of species is still uncertain. While there are more than 300 published species names, the actual number could be as low as 100-150 species.

Laminaria

Laminaria is a genus of 31 species of brown algae commonly called "kelp". Some species are also referred to as tangle. This economically important genus is characterized by long, leathery laminae and relatively large size. Some species are referred to by the common name Devil's apron, due to their shape, or sea colander, due to the perforations present on the lamina. It is found in the north Atlantic Ocean and the northern Pacific Ocean at depths from 8 to 30 m (26 to 98 ft) (exceptionally to 120 m (390 ft) in the warmer waters of the Mediterranean Sea and off Brazil). Laminaria form a habitat for many fish and invertebrates. The life cycle of Laminaria has heteromorphic alternation of generations which differs from Fucus. At meiosis the male and female zoospores are produced separately, then germinate into male and female gametophytes. The female egg matures in the oogonium until the male sperm fertilizes it. Life-Cycle: The most apparent form of Laminaria is its sporophyte phase, a structure composed of the holdfast, the stipe, and the blades. While it spends its time predominately in the sporophyte phase, it alternates between the sporophyte and its microscopic gametophyte phase.Laminaria japonica (J. E. Areschoug – Japón) is now regarded as a synonym of Saccharina japonica and Laminaria saccharina is now classified as Saccharina latissima.

Lycopodiophyta

The Division Lycopodiophyta (sometimes called lycophyta or lycopods) is a tracheophyte subgroup of the Kingdom Plantae. It is one of the oldest lineages of extant (living) vascular plants and contains extinct plants like Baragwanathia that have been dated from the Silurian (ca. 425 million years ago). Members of Lycopodiophyta were some of the dominating plant species of the Carboniferous period. These species reproduce by shedding spores and have macroscopic alternation of generations, although some are homosporous while others are heterosporous. Most members of Lycopodiophyta bear a protostele, and the sporophyte generation is dominant. They differ from all other vascular plants in having microphylls, leaves that have only a single vascular trace (vein) rather than the much more complex megaphylls found in ferns and seed plants.

Marchantiophyta

The Marchantiophyta (listen) are a division of non-vascular land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of genetic information.

It is estimated that there are about 9000 species of liverworts. Some of the more familiar species grow as a flattened leafless thallus, but most species are leafy with a form very much like a flattened moss. Leafy species can be distinguished from the apparently similar mosses on the basis of a number of features, including their single-celled rhizoids. Leafy liverworts also differ from most (but not all) mosses in that their leaves never have a costa (present in many mosses) and may bear marginal cilia (very rare in mosses). Other differences are not universal for all mosses and liverworts, but the occurrence of leaves arranged in three ranks, the presence of deep lobes or segmented leaves, or a lack of clearly differentiated stem and leaves all point to the plant being a liverwort.

Liverworts are typically small, usually from 2–20 mm wide with individual plants less than 10 cm long, and are therefore often overlooked. However, certain species may cover large patches of ground, rocks, trees or any other reasonably firm substrate on which they occur. They are distributed globally in almost every available habitat, most often in humid locations although there are desert and Arctic species as well. Some species can be a nuisance in shady greenhouses or a weed in gardens.

Moss

Mosses are small flowerless plants that typically form dense green clumps or mats, often in damp or shady locations. The individual plants are usually composed of simple leaves that are generally only one cell thick, attached to a stem that may be branched or unbranched and has only a limited role in conducting water and nutrients. Although some species have conducting tissues, these are generally poorly developed and structurally different from similar tissue found in vascular plants. Mosses do not have seeds and after fertilisation develop sporophytes with unbranched stalks topped with single capsules containing spores. They are typically 0.2–10 cm (0.1–3.9 in) tall, though some species are much larger. Dawsonia, the tallest moss in the world, can grow to 50 cm (20 in) in height.

Mosses are commonly confused with lichens, hornworts, and liverworts. Lichens may superficially resemble mosses, and sometimes have common names that include the word "moss" (e.g., "reindeer moss" or "Iceland moss"), but they are not related to mosses. Mosses were formerly grouped with the hornworts and liverworts as "non-vascular" plants in the division "bryophytes", all of them having the haploid gametophyte generation as the dominant phase of the life cycle. This contrasts with the pattern in all vascular plants (seed plants and pteridophytes), where the diploid sporophyte generation is dominant.

Mosses are now classified on their own as the division Bryophyta. There are approximately 12,000 species.The main commercial significance of mosses is as the main constituent of peat (mostly the genus Sphagnum), although they are also used for decorative purposes, such as in gardens and in the florist trade. Traditional uses of mosses included as insulation and for the ability to absorb liquids up to 20 times their weight.

Non-vascular plant

Non-vascular plants are plants without a vascular system consisting of xylem and phloem. Although non-vascular plants lack these particular tissues, many possess simpler tissues that are specialized for internal transport of water.

Non-vascular plants include two distantly related groups:

Bryophytes, an informal group that is now treated as three separate land plant Divisions, namely Bryophyta (mosses), Marchantiophyta (liverworts), and Anthocerotophyta (hornworts). In all bryophytes, the primary plants are the haploid gametophytes, with the only diploid portion being the attached sporophyte, consisting of a stalk and sporangium. Because these plants lack lignified water-conducting tissues, they can't become as tall as most vascular plants.

Algae - especially the green algae. Recent studies have demonstrated that the algae consist of several unrelated groups. It turns out that the common features of living in water and photosynthesis were misleading as indicators of close relationship. Only those groups of algae included in the Viridiplantae are still considered relatives of land plants.These groups are sometimes referred to as "lower plants", referring to their status as the earliest plant groups to evolve, but the usage is imprecise, since both groups are polyphyletic and may be used to include vascular cryptogams, such as the ferns and fern allies that reproduce using spores. Non-vascular plants are often among the first species to move into new and inhospitable territories, along with prokaryotes and protists, and thus function as pioneer species.

Non-vascular plants do not have a wide variety of specialized tissue types. Mosses and leafy liverworts have structures called phyllids that look like leaves, but are not true leaves because they are single sheets of cells with no internal air spaces, no cuticle or stomata and no xylem or phloem. Consequently, phyllids are unable to control the rate of water loss from their tissues and are said to be poikilohydric. Some liverworts, such as Marchantia have a cuticle and the sporophytes of mosses have both cuticles and stomata, which were important in the evolution of land plants.All land plants have a life cycle with an alternation of generations between a diploid sporophyte and a haploid gametophyte, but in all non-vascular land plants the gametophyte generation is dominant. In these plants, the sporophytes grow from and are dependent on gametophytes for taking in water and mineral nutrients and for provision of photosynthate, the products of photosynthesis.

Perianth

The perianth (perigonium, perigon or perigone) is the non-reproductive part of the flower, and structure that forms an envelope surrounding the sexual organs, consisting of the calyx (sepals) and the corolla (petals). The term perianth is derived from the Greek περί, peri, meaning around, and άνθος, anthos, meaning flower, while perigonium is derived from gonos, meaning seed, i.e. sexual organs.

In the mosses and liverworts (Marchantiophyta), the perianth is the sterile tubelike tissue that surrounds the female reproductive structure (or developing sporophyte).

Plant reproduction

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 the parent or parents. Asexual reproduction produces new individuals without the fusion of gametes, genetically identical to the parent plants and each other, except when mutations occur. In seed plants, the offspring can be packaged in a protective seed, which is used as an agent of dispersal.

Polysporangiophyte

Polysporangiophytes, also called polysporangiates or formally Polysporangiophyta, are plants in which the spore-bearing generation (sporophyte) has branching stems (axes) that terminate in sporangia. The name literally means many sporangia plant. The clade includes all land plants (embryophytes) except for the bryophytes (liverworts, mosses and hornworts) whose sporophytes are normally unbranched, even if a few exceptional cases occur. While the definition is independent of the presence of vascular tissue, all living polysporangiophytes also have vascular tissue, i.e., are vascular plants or tracheophytes. Fossil polysporangiophytes are known that have no vascular tissue, and so are not tracheophytes.

Pteridophyte

A pteridophyte is a vascular plant (with xylem and phloem) that reproduces using spores. Because pteridophytes produce neither flowers nor seeds, they are also referred to as "cryptogams", meaning that their means of reproduction is hidden. The pteridophytes include the ferns, horsetails, and the lycophytes (clubmosses, spikemosses, and quillworts). These are not a monophyletic group because ferns and horsetails are more closely related to seed plants than to the lycophytes. Therefore, "Pteridophyta" is no longer a widely accepted taxon, although the term pteridophyte remains in common parlance, as do pteridology and pteridologist as a science and its practitioner, to indicate lycophytes and ferns as an informal grouping, such as the International Association of Pteridologists and the Pteridophyte Phylogeny Group.

Sorus

A sorus (pl. sori) is a cluster of sporangia (structures producing and containing spores) in ferns and fungi. This New Latin word is from Ancient Greek σωρός (sōrós 'stack, pile, heap').

In lichens and other fungi, the sorus is surrounded by an external layer. In some red algae it may take the form of a depression into the thallus.

In ferns, these form a yellowish or brownish mass on the edge or underside of a fertile frond. In some species, they are protected during development by a scale or film of tissue called the indusium, which forms an umbrella-like cover.

Sori occur on the sporophyte generation, the sporangia within producing haploid meiospores. As the sporongia mature, the indusium shrivels so that spore release is unimpeded. The sporangia then burst and release the spores.

The shape, arrangement, and location of the sori are often valuable clues in the identification of fern taxa. Sori may be circular or linear. They may be arranged in rows, either parallel or oblique to the costa, or randomly. Their location may be marginal or set away from the margin on the frond lamina. The presence or absence of indusium is also used to identify fern taxa.

Spore

In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, fungi and protozoa. Bacterial spores are not part of a sexual cycle but are resistant structures used for survival under unfavourable conditions. Myxozoan spores release 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.Spores are usually haploid and unicellular and are produced by meiosis in the sporangium of a diploid sporophyte. 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.

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