Contents 1 Origin of endosperm 1.1 Double fertilization 1.2 Endosperm formation 2 Evolutionary origins 3 The role of endosperm in seed development 4 Cereal grains 5 See also 6 References 7 External links

Origin of endosperm[edit] Ancestral flowering plants have seeds with small embryos and abundant endosperm, and the evolutionary development of flowering plants tends to show a trend towards plants with mature seeds with little or no endosperm. In more derived flowering plants the embryo occupies most of the seed and the endosperm is non developed or consumed before the seed matures.[2][3] Double fertilization[edit] Endosperm is formed when the two sperm nuclei inside a pollen grain reach the interior of a female gametophyte (sometimes called the embryo sac). One sperm nucleus fertilizes the egg cell, forming a zygote, while the other sperm nucleus usually fuses with the binucleate central cell, forming a primary endosperm cell (its nucleus is often called the triple fusion nucleus). That cell created in the process of double fertilization develops into the endosperm. Because it is formed by a separate fertilization, the endosperm constitutes an organism separate from the growing embryo. About 70% of angiosperm species have endosperm cells that are polyploid.[4] These are typically triploid (containing three sets of chromosomes), but can vary widely from diploid (2n) to 15n.[5] One species of flowering plant, Nuphar polysepala, has been shown to have endosperm that is diploid, resulting from the fusion of a pollen nucleus with one, rather than two, maternal nuclei. The same is supposed for some other basal angiosperms.[6] It is believed that early in the development of angiosperm lineages, there was a duplication in this mode of reproduction, producing seven-celled/eight-nucleate female gametophytes, and triploid endosperms with a 2:1 maternal to paternal genome ratio.[7] Endosperm formation[edit] There are three types of Endosperm development: Nuclear endosperm formation - where repeated free-nuclear divisions take place; if a cell wall is formed it will form after free-nuclear divisions. Commonly referred to as liquid endosperm. Coconut water is an example of this. Cellular endosperm formation - where a cell-wall formation is coincident with nuclear divisions. Coconut meat is cellular endosperm. Acoraceae has cellular endosperm development while other monocots are helobial. Helobial endosperm formation - Where a cell wall is laid down between the first two nuclei, after which one half develops endosperm along the cellular pattern and the other half along the nuclear pattern.

Evolutionary origins[edit] The evolutionary origins of double fertilization and endosperm are unclear, attracting researcher attention for over a century. There are the two major hypotheses:[8] The double fertilization initially used to produce two identical, independent embryos ("twins"). Later these embryos acquired different roles, one growing into the mature organism, and another merely supporting it. Thus the early endosperm was probably diploid, like the embryo. Some gymnosperms, such as Ephedra (genus), may produce twin embryos by double fertilization. Either of these two embryos is capable of filling in the seed, but normally only one develops further (the other eventually aborts). Also, most basal angiosperms still contain the four-cell embryo sac and produce diploid endosperms. Endosperm is the evolutionary remnant of the actual gametophyte, similar to the complex multicellular gametophytes found in gymnosperms. In this case, acquisition of the additional nucleus from the sperm cell is a later evolutionary step. This nucleus may provide the parental (not only maternal) organism with some control over endosperm development. Becoming triploid or polyploid are later evolutionary steps of this "primary gametophyte". Nonflowering seed plants (conifers, cycads, Ginkgo, Ephedra) form a large homozygous female gametophyte to nourish the embryo within a seed.[9]

The role of endosperm in seed development[edit] In some groups (e.g. grains of the family Poaceae) the endosperm persists to the mature seed stage as a storage tissue, in which case the seeds are called "albuminous" or "endospermous", and in others it is absorbed during embryo development (e.g., most members of the family Fabaceae, including the common bean, Phaseolus vulgaris), in which case the seeds are called "exalbuminous" or "cotyledonous" and the function of storage tissue is performed by enlarged cotyledons ("seed leaves"). In certain species (e.g. corn, Zea mays); the storage function is distributed between both endosperm and the embryo. Some mature endosperm tissue stores fats (e.g. castor bean, Ricinus communis) and others (including grains, such as wheat and corn) store mainly starches. The dust-like seeds of orchids have no endosperm. Orchid seedlings are mycoheterotrophic in their early development. In some other species, such as coffee, the endosperm also does not develop.[10] Instead, the nucellus produces a nutritive tissue termed "perisperm". The endosperm of some species is responsible for seed dormancy.[11] Endosperm tissue also mediates the transfer of nutrients from the mother plant to the embryo, it acts as a location for gene imprinting, and is responsible for aborting seeds produced from genetically mismatched parents.[4] In angiosperms, the endosperm contain hormones such as cytokinins, which regulate cellular differentiation and embryonic organ formation.[12]

Cereal grains[edit] Cereal crops are grown for their edible fruit (grains or caryopses), which are primarily endosperm. In the caryopsis, the thin fruit wall is fused to the seed coat. Therefore, the nutritious part of the grain is the seed and its endosperm. In some cases (e.g. wheat, rice) the endosperm is selectively retained in food processing (commonly called white flour), and the embryo (germ) and seed coat (bran) removed. The processed grain has a lower quality of nutrition. Endosperm thus has an important role within the human diet worldwide. The aleurone is the outer layer of endosperm cells, present in all small grains and retained in many dicots with transient endosperm. The cereal aleurone functions for both storage and digestion. During germination, it secretes the amylase enzyme that breaks down endosperm starch into sugars to nourish the growing seedling.[13] [14]

See also[edit] Seed Ovule

References[edit] ^ retrieved 14 July 2010 ^ "The Seed Biology Place - Seed Dormancy". Retrieved 2014-02-05.  ^ Friedman, William E. (1998), "The evolution of double fertilization and endosperm: an "historical" perspective", Sexual Plant Reproduction, 11: 6, doi:10.1007/s004970050114  ^ a b Olsen, By Odd-Arne (2007), "Endosperm: Developmental and Molecular Biology", ISBN 3-540-71234-8  ^ "Figure 1". doi:10.1186/gb-2002-3-9-reviews1026. Retrieved 2014-02-05.  ^ Williams J. H., Friedman W. E. (2002). "Identification of diploid endosperm in an early angiosperm lineage". Letters to Nature. doi:10.1038/415522a.  ^ Friedman, William E.; Williams, JH (2003), "Modularity of the Angiosperm Female Gametophyte and Its Bearing on the Early Evolution of Endosperm in Flowering Plants", Evolution, 57 (2): 216–30, doi:10.1111/j.0014-3820.2003.tb00257.x, PMID 12683519  ^ Célia Baroux, Charles Spillane, Ueli Grossniklaus (2002). Evolutionary origins of the endosperm in flowering plants. Genome Biol. 3(9) ^ [Friedman WE (1995)] Organismal duplication, inclusive fitness theory, and altruism: understanding the evolution of endosperm and the angiosperm reproductive syndrome. Proc Natl Acad Sci U S A. 1995 Apr 25;92(9):3913-7.PMID 11607532pdf ^ Houk, W. G. (1938). "Endosperm and Perisperm of Coffee with Notes on the Morphology of the Ovule and Seed Development". American Journal of Botany. 25 (1): 56–61. doi:10.2307/2436631.  ^ Mechanisms of plant growth and improved productivity: modern approaches edited by Amarjit S. Basra 1994. ISBN 0-8247-9192-4 p. ^ Pearson, Lorentz C. (1995), The diversity and evolution of plants, Boca Raton: CRC Press, p. 547, ISBN 0-8493-2483-1  ^ Becraft, P.W., and Yi, G. (2011). Regulation of aleurone development in cereal grains. J Exp Bot 62, 1669-1675. ^ Becraft, P.W., and Gutierrez-Marcos, J. (2012). Endosperm development: dynamic processes and cellular innovations underlying sibling altruism. WIREs Dev Biol 1, 579-593.

External links[edit]  Beach, Chandler B., ed. (1914). "Endosperm". The New Student's Reference Work. Chicago: F. E. Compton and Co.  Endosperm: the pivot of the sexual conflict in flowering plants at Earthling Nature v t e Botany History of botany Subdisciplines Plant systematics Ethnobotany Paleobotany Plant anatomy Plant ecology Phytogeography Geobotany Flora Phytochemistry Plant pathology Bryology Phycology Floristics Dendrology Plant groups Algae Archaeplastida Bryophyte Non-vascular plants Vascular plants Spermatophytes Pteridophyte Gymnosperm Angiosperm Plant morphology (glossary) Plant cells Cell wall Phragmoplast Plastid Plasmodesma Vacuole Tissues Meristem Vascular tissue Vascular bundle Ground tissue Mesophyll Cork Wood Storage organs Vegetative Root Rhizoid Bulb Rhizome Shoot Stem Leaf Petiole Cataphyll Bud Sessility Reproductive (Flower) Flower development Inflorescence Umbel Raceme Bract Pedicellate Flower Whorl Floral symmetry Floral diagram Floral formula Receptacle Hypanthium (Floral cup) Perianth Tepal Petal Sepal Sporophyll Gynoecium Ovary Ovule Stigma Archegonium Androecium Stamen Staminode Pollen Tapetum Gynandrium Gametophyte Sporophyte Plant embryo Fruit Fruit anatomy Berry Capsule Seed Seed dispersal Endosperm Surface structures Epicuticular wax Plant cuticle Epidermis Stoma Nectary Trichome Prickle Plant physiology Materials Nutrition Photosynthesis Chlorophyll Plant hormone Transpiration Turgor pressure Bulk flow Aleurone Phytomelanin Sugar Sap Starch Cellulose Plant growth and habit Secondary growth Woody plants Herbaceous plants Habit Vines Lianas Shrubs Subshrubs Trees Succulent plants Reproduction Evolution Ecology Alternation of generations Sporangium Spore Microsporangia Microspore Megasporangium Megaspore Pollination Pollinators Pollen tube Double fertilization Germination Evolutionary development Evolutionary history timeline Hardiness zone Plant taxonomy History of plant systematics Herbarium Biological classification Botanical nomenclature Botanical name Correct name Author citation International Code of Nomenclature for algae, fungi, and plants (ICN) - for Cultivated Plants (ICNCP) Taxonomic rank International Association for Plant Taxonomy (IAPT) Plant taxonomy systems Cultivated plant taxonomy Citrus taxonomy cultigen cultivar Group grex Practice Agronomy Floriculture Forestry Horticulture Lists Related topics Botanical terms Botanists by author abbreviation Botanical expedition Category Portal WikiProject Authority control GND: 4367419-7 Retrieved from "" Categories: Plant morphologyPlant physiologyHidden categories: Wikipedia articles incorporating citation to the NSRWWikipedia articles incorporating citation to the NSRW with an wstitle parameterWikipedia articles with GND identifiers

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