Megaspore

Megaspores, also called macrospores, are a type of spore that is present in heterosporous plants. These plants have two spore types, megaspores and microspores. Generally speaking, the megaspore, or large spore, germinates into a female gametophyte, which produces egg cells. These are fertilized by sperm produced by the male gametophyte developing from the microspore. Heterosporous plants include the following:

Selaginella heterospores.jpeg
Microscopic photo of spores (in red) of Selaginella. The large three spores at the top are megaspores whereas the numerous smaller red spores at the bottom are microspores.

Megasporogenesis

In gymnosperms and flowering plants, the megaspore is produced inside the nucleus of the ovule. During megasporogenesis, a diploid precursor cell, the megasporocyte or megaspore mother cell, undergoes meiosis to produce initially four haploid cells (the megaspores).[1] Angiosperms exhibit three patterns of megasporogenesis: monosporic, bisporic, and tetrasporic, also known as the Polygonum type, the Alisma type, and the Drusa type, respectively. The monosporic pattern occurs most frequently (>70% of angiosperms) and is found in many economically and biologically important groups such as Brassicaceae (e.g., Arabidopsis, Capsella, Brassica), Gramineae (e.g., maize, rice, wheat), Malvaceae (e.g., cotton), Leguminoseae (e.g., beans, soybean), and Solanaceae (e.g., pepper, tobacco, tomato, potato, petunia).[2]

This pattern is characterized by cell plate formation after meiosis 1 & 2, which results in four one-nucleate megaspores, of which three degenerate. The bisporic pattern is characterized by cell plate formation only after meiosis 1, and results in two two-nucleate megaspores, of which one degenerates. The tetrasporic pattern is characterized by cell plates failing to form after either meiosis 1 or 2, and results in one four-nucleate megaspore. Therefore, each pattern gives rise to a single functional megaspore which contains one, two, or four meiotic nuclei, respectively.[2] The megaspore then undergoes megagametogenesis to give rise to the female [gametophyte].

Megagametogenesis

Ovule-Gymno-Angio-en
Plant ovules with megasporocytes before meiosis: Gymnosperm ovule on left, angiosperm ovule (inside ovary) on right

After megasporogenesis, the megaspore develops into the female gametophyte (the embryo sac) in a process called megagametogenesis. The process of megagametogenesis varies depending on which pattern of megasporogenesis occurred. Some species, such as Tridax trilobata, Ehretia laevis, and Alectra thomsoni, can undergo different patterns of megasporogenesis and therefore different patterns of megagametogenesis. If the monosporic pattern occurred, the single nucleus undergoes mitosis three times, producing an eight-nucleate cell. These eight nuclei are arranged into two groups of four. These groups both send a nucleus to the center of the cell; these become the polar nuclei. Depending on the species, these nuclei fuse together before or upon fertilization of the central cell. The three nuclei at the end of the cell near the micropylar become the egg apparatus, with an egg cell in the center and two synergids. At the other end of the cell, a cell wall forms around the nuclei and forms the antipodals. Therefore, the resulting embryo sac is a seven-celled structure consisting of one central cell, one egg cell, two synergid cells, and three antipodal cells.[2][3]

The bisporic and tetrasporic patterns undergo varying processes and result in varying embryo sacs as well. In Lilium which has a tetrasporic pattern, the central cell of the embryo sac is 4N. Therefore, upon fertilization the endosperm will be 5N rather than the typical 3N.[4]

See also

References

  1. ^ Estrada-Luna, A. A.; W. Huanca-Mamani; G. Acosta-García; G. León-Martínez; A. Becerra-Flora; R. Pérez-Ruíz; J. -Ph. Vielle-Calzada (Mar–Apr 2002). "Beyond Promiscuity: From Sexuality to Apomixis in Flowering Plants". In Vitro Cellular & Developmental Biology – Plant. 38 (2): 146–151. doi:10.1079/ivp2001278. JSTOR 20065024.
  2. ^ a b c Yadegaria, Ramin; Gary N. Drewsb (2004-04-09). "Female Gametophyte Development". The Plant Cell. 16 (Suppl): S133–S141, Supplement. doi:10.1105/tpc.018192. PMC 2643389. Retrieved 2010-11-28.
  3. ^ Raven, Peter H.; Evert, Ray Franklin.; Eichhorn, Susan E. (2005). Biology of Plants. New York, NY: W.H. Freeman. ISBN 0-7167-1007-2.
  4. ^ Sargent, E. (1900). "Recent work on the results of fertilization in angiosperms". Annals of Botany. 14: 689–712. Archived from the original on 2010-12-10. Retrieved 2010-11-28.
Abies balsamea

Abies balsamea or balsam fir is a North American fir, native to most of eastern and central Canada (Newfoundland west to central Alberta) and the northeastern United States (Minnesota east to Maine, and south in the Appalachian Mountains to West Virginia).It is celebrated for its rich green needles, natural conical shape, and needle retention after being cut, and it is notably the most fragrant of all Christmas tree varieties.

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 haploid sexual and diploid asexual 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.

Azolla

Azolla (mosquito fern, duckweed fern, fairy moss, water fern) is a genus of seven species of aquatic ferns in the family Salviniaceae. They are extremely reduced in form and specialized, looking nothing like other typical ferns but more resembling duckweed or some mosses. Azolla filiculoides is one of just two fern species for which a reference genome has been published.

Double fertilization

Double fertilization is a complex fertilization mechanism of flowering plants (angiosperms). This process involves the joining of a female gametophyte (megagametophyte, also called the embryo sac) 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.

The cells of an unfertilized ovule are 8 in number and arranged in the form of 3+2+3 (from top to bottom) i.e. 3 antipodal cells, 2 polar central cells, 2 synergids & 1 egg cell. One sperm fertilizes the egg cell and the other sperm combines with the two polar nuclei of the large central cell of the megagametophyte. The haploid sperm and haploid egg combine to form a diploid zygote,the process being called syngamy, while the other sperm and the two haploid polar nuclei of the large central cell of the megagametophyte form a triploid nucleus (triple fusion). Some plants may form polyploid nuclei. The large cell of the gametophyte will then develop into the endosperm, a nutrient-rich tissue which provides nourishment to the developing embryo. The ovary, surrounding the ovules, develops into the fruit, which protects the seeds and may function to disperse them.The two central cell maternal nuclei (polar nuclei) that contribute to the endosperm, arise by mitosis from the same single meiotic product that gave rise to the egg. The maternal contribution to the genetic constitution of the triploid endosperm is double that of the embryo.

In a study conducted in 2008 of the plant Arabidopsis thaliana, the migration of male nuclei inside the female gamete, in fusion with the female nuclei, has been documented for the first time using in vivo imaging. Some of the genes involved in the migration and fusion process have also been determined.Evidence of double fertilization in Gnetales, which are non-flowering seed plants, has been reported.

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.

Heterospory

Heterospory is the production of spores of two different sizes and sexes by the sporophytes of land plants. The smaller of these, the microspore, is male and the larger megaspore is female. Heterospory evolved during the Devonian period from isospory independently in several plant groups: the clubmosses, the arborescent horsetails, and progymnosperms. This occurred as part of the process of evolution of the timing of sex differentiation.

Isoetes acadiensis

Acadian quillwort (Isoetes acadiensis) is a species of quillwort in the Isoetaceae family described by Kott in 1981. It can be found along the shores of lakes, ponds, and rivers in Newfoundland, Nova Scotia, and New Brunswick, as well as in the American states Maine, Massachusetts, and New Hampshire. It has a similar distribution to that of I. tuckermanii. It bears 9 to 35 mostly recurved leaves, each 5–21 cm long. The leaves are usually dark green, though can occasionally be tinged with red. The sporangium can be up to five millimeters long and 3 millimeters in length, covered one sixth to one third by the velum. The spherical megaspores are 400-570 micrometers in diameter, and bear smooth ridges. The kidney shaped microspores are 25 to 30 micrometers long. It was originally believed to be a member of Isoetes hieroglyphica because of their similar megaspore structure.

Isoetes tuckermanii

Isoetes tuckermanii (Tuckerman's quillwort) is a tetraploid species of quillwort in the Isoetaceae family. It can be found in shallow water in Newfoundland, Nova Scotia, New Brunswick, and south through the New England states to Maryland. It bears 10 to 45 long bright green to yellow green leaves that are 4 to 25 centimeters long, usually erect, but sometimes recurved. The velum covers one fourth or less of the sporangium, which is usually unspotted, 5 millimeters long, and 3 millimeters wide. The white spherical megaspores are 400 to 650 micrometers in diameter, and bear rough-crested ridges that form a hexagonal honeycomb shape. The kidney shaped microspores are 24 to 33 micrometers long, bearing tubercles. It is very similar to I. macrospora, only reliably distinguishable by cytology or through careful megaspore measurement.

Lyginopteridales

The Lyginopteridales were the archetypal pteridosperms: They were the first plant fossils to be described as pteridosperms and, thus, the group on which the concept of pteridosperms was first developed; they are the stratigraphically oldest-known pteridosperms, occurring first in late Devonian strata; and they have the most primitive features, most notably in the structure of their ovules. They probably evolved from a group of Late Devonian progymnosperms known as the Aneurophytales, which had large, compound frond-like leaves. The Lyginopteridales became the most abundant group of pteridosperms during Mississippian times, and included both trees and smaller plants. During early and most of middle Pennsylvanian times the Medullosales took over as the more important of the larger pteridosperms but the Lyginopteridales continued to flourish as climbing (lianesent) and scrambling plants. However, later in Middle Pennsylvanian times the Lyginopteridales went into serious decline, probably being out-competed by the Callistophytales that occupied similar ecological niches but had more sophisticated reproductive strategies. A few species continued into Late Pennsylvanian times, and in China persisted into Early Permian (Asselian) times, but then became extinct. Most evidence of the Lyginopteridales suggests that they grew in tropical latitudes of the time, in North America, Europe and China.

Megagametogenesis

Megagametogenesis is the process of maturation of the female gametophyte, or megagametophyte, in plants During the process of megagametogenesis, the megaspore, which arises from megasporogenesis, develops into the embryo sac, which is where the female gamete is housed. These megaspores then develop into the haploid female gametophytes. This occurs within the ovule, which is housed inside the ovary.

Megaspore mother cell

A megaspore mother cell, or megasporocyte, is a diploid cell in plants in which meiosis will occur, resulting in the production of four haploid megaspores. At least one of the spores develop into haploid female gametophytes (megagametophytes). The megaspore mother cell arises within the megasporangium tissue.

In flowering plants the megasporangium is also called the nucellus, and the female gametophyte is sometimes called the embryo sac.

Microspore

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.

Ovule

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 (or remnant of the megasporangium), and the female gametophyte (formed from a haploid megaspore) 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.

Raceme

A raceme ( or ) is an unbranched, indeterminate type of inflorescence bearing pedicellate flowers (flowers having short floral stalks called pedicels) along its axis. In botany, an axis means a shoot, in this case one bearing the flowers. In indeterminate inflorescence-like racemes, the oldest flowers are borne towards the base and new flowers are produced as the shoot grows, with no predetermined growth limit. A plant that flowers on a showy raceme may have this reflected in its scientific name, e.g. Cimicifuga racemosa. A compound raceme, also called a panicle, has a branching main axis. Examples of racemes occur on mustard (genus Brassica) and radish (genus Raphanus) plants.

Salviniales

The order Salviniales (formerly known as the Hydropteridales and including the former Marsileales) is an order of ferns in the class Polypodiopsida.

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.

Sporocarp (ferns)

A sporocarp is a specialised type of structure found in some ferns whose primary function is the production and release of spores.

Among ferns, sporocarps are found only in the three families of the order Salviniales, a group that is both aquatic and heterosporous. The term actually refers to two very different structures. In the Azollaceae and Salviniaceae, the sporocarp is nothing more than a modified sorus, a single cluster of spore-producing tissues enclosed by a thin sphere of tissue and attached to the leaves.

In the Marsileaceae (water-clover family), however, the sporocarp is a more elaborate structure formed from an entire leaf whose development and form is greatly modified. These are hairy, short-stalked, bean-shaped structures (usually 3 to 8 mm in diameter) with a hardened outer covering. This outer covering is tough and resistant to drying out, allowing the spores inside to survive unfavorable conditions such as winter frost or summer desiccation. Despite this toughness, the sporocarps will open readily in water if conditions are favorable, and specimens have been successfully germinated after being stored for more than forty years. Each growing season, only one sporocarp develops per node along the rhizome near the base of the other leaf-stalks.

The sporocarps are functionally and developmentally modified leaves, although they have much shorter stalks than the vegetative leaves. Inside the sporocarp, the modified leaflets bear several sori, each of which consists of several sporangia covered by a thin hood of tissue (the indusium). Each sorus includes a mix of two types of sporangium, each type producing only one of two kinds of spores. Toward the center of each sorus and developing first are the megasporangia, each of which will produce a single large female megaspore. Surrounding them at the edge of the sorus and developing later are the microsporangia, each of which will produce many small male microspores.

Sporogenesis

Sporogenesis is the production of spores in biology. The term is also used to refer to the process of reproduction via spores. Reproductive spores were found to be formed in eukaryotic organisms, such as plants, algae and fungi, during their normal reproductive life cycle. Dormant spores are formed, for example by certain fungi and algae, primarily in response to unfavorable growing conditions. Most eukaryotic spores are haploid and form through cell division, though some types are diploid or dikaryons and form through cell fusion.

Sporophyte

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, (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.

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.

Because this embryo-nurturing feature of the life cycle is common to all land plants they are known collectively as the embryophytes.

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.

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