Flowering plant
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. They include all forbs (flowering plants without a woody stem), grasses and grass-like plants, a vast majority of broad-leaved trees, shrubs and vines, and most aquatic plants. The term "angiosperm" is derived from the Greek words ἀγγεῖον /  ('container, vessel') and σπέρμα /  ('seed'), meaning that the seeds are enclosed within a fruit. They are by far 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 ().

Angiosperms are distinguished from the other seed-producing plants, the gymnosperms, by having flowers, xylem consisting of vessel elements instead of tracheids, endosperm within their seeds, and fruits that completely envelop the seeds.
The ancestors of flowering plants diverged from the common ancestor of all living gymnosperms before the end of the Carboniferous, over 300 million years ago. In the Cretaceous, angiosperms diversified explosively, becoming the dominant group of plants across the planet.

Agriculture is almost entirely dependent on angiosperms, and a small number of flowering plant families supply nearly all plant-based food and livestock feed. Rice, maize, and wheat provide half of the world's calorie intake, and all three plants are cereals from the Poaceae family (colloquially known as grasses). Other families provide materials such as wood, paper and cotton, and supply numerous ingredients for traditional and modern medicines. Flowering plants are also commonly grown for decorative purposes, with certain flowers playing a significant role in many cultures.

Out of the "Big Five" extinction events in Earth's history, only the Cretaceous–Paleogene extinction event had occurred while angiosperms dominated plant life on the planet. Today, the Holocene extinction affects all kingdoms of complex life on Earth, and conservation measures are necessary to protect plants in their habitats in the wild (in situ), or failing that, ex situ in seed banks or artificial habitats like botanic gardens. Otherwise, around 40% of plant species may become extinct due to human actions such as habitat destruction, introduction of invasive species, unsustainable logging and collection of medicinal or ornamental plants. Further, climate change is starting to impact plants and is likely to cause many species to become extinct by 2100.

 Distinguishing features 

Angiosperms are terrestrial vascular plants; like the gymnosperms, they have roots, stems, leaves, and seeds. They differ from other seed plants in several ways.

 Feature  Description  ImageFlowers The reproductive organs of flowering plants, not found in any other seed plants.  thumb|none|A Narcissus flower in section. Petals and sepals are replaced here by a fused tube, the corona, and tepals.Reduced gametophytes, three cells in male, seven cells with eight nuclei in female The gametophytes are smaller than those of gymnosperms. The smaller size of the pollen reduces the time between pollination and fertilization, which in gymnosperms is up to a year.   thumb|none|Embryo sac is a reduced female gametophyte.Endosperm Endosperm forms after fertilization but before the zygote divides. It provides food for the developing embryo, the cotyledons, and sometimes the seedling. Closed carpel enclosing the ovules. Once the ovules are fertilised, the carpels, often with surrounding tissues, develop into fruits. Gymnosperms have unenclosed seeds.  thumb|none|Peas (seeds, from ovules) inside pod (fruit, from fertilised carpel). Xylem made of vessel elements  Open vessel elements are stacked end to end to form continuous tubes, whereas gymnosperm xylem is made of tapered tracheids connected by small pits.  thumb|none| Xylem vessels (long tubes).

 Diversity 

 Ecological diversity 

The largest angiosperms are Eucalyptus gum trees of Australia, and Shorea faguetiana, dipterocarp rainforest trees of Southeast Asia, both of which can reach almost  in height. The smallest are Wolffia duckweeds which float on freshwater, each plant less than  across.

Considering their method of obtaining energy, some 99% of flowering plants are photosynthetic autotrophs, deriving their energy from sunlight and using it to create molecules such as sugars. The remainder are parasitic, whether on fungi like the orchids for part or all of their life-cycle, or on other plants, either wholly like the broomrapes, Orobanche, or partially like the witchweeds, Striga.

In terms of their environment, flowering plants are cosmopolitan, occupying a wide range of habitats on land, in fresh water and in the sea. On land, they are the dominant plant group in every habitat except for frigid moss-lichen tundra and coniferous forest. The seagrasses in the Alismatales grow in marine environments, spreading with rhizomes that grow through the mud in sheltered coastal waters.

Some specialised angiosperms are able to flourish in extremely acid or alkaline habitats. The sundews, many of which live in nutrient-poor acid bogs, are carnivorous plants, able to derive nutrients such as nitrate from the bodies of trapped insects. Other flowers such as Gentiana verna, the spring gentian, are adapted to the alkaline conditions found on calcium-rich chalk and limestone, which give rise to often dry topographies such as limestone pavement.

As for their growth habit, the flowering plants range from small, soft herbaceous plants, often living as annuals or biennials that set seed and die after one growing season, to large perennial woody trees that may live for many centuries and grow to many metres in height. Some species grow tall without being self-supporting like trees by climbing on other plants in the manner of vines or lianas.

 Taxonomic diversity 

The number of species of flowering plants is estimated to be in the range of 250,000 to 400,000. This compares to around 12,000 species of moss and 11,000 species of pteridophytes.  The APG system seeks to determine the number of families, mostly by molecular phylogenetics. In the 2009 APG III there were 415 families. The 2016 APG IV added five new orders (Boraginales, Dilleniales, Icacinales, Metteniusales and Vahliales), along with some new families, making a total of 64 angiosperm orders and 416 families.

The diversity of flowering plants is not evenly distributed. Nearly all species belong to the eudicot (75%), monocot (23%), and magnoliid (2%) clades. The remaining five clades contain a little over 250 species in total; i.e. less than 0.1% of flowering plant diversity, divided among nine families. The 25 most species-rich of 443 families,  containing over 166,000 species between them in their APG circumscriptions, are:

+ The 25 largest angiosperm families Group  Family  English name  No. of spp.  Eudicot   Asteraceae or Compositae   daisy   22,750  Monocot   Orchidaceae   orchid   21,950  Eudicot   Fabaceae or Leguminosae   pea, legume   19,400  Eudicot   Rubiaceae   madder   13,150  Monocot   Poaceae or Gramineae   grass   10,035  Eudicot   Lamiaceae or Labiatae   mint   7,175  Eudicot   Euphorbiaceae   spurge   5,735  Eudicot   Melastomataceae   melastome   5,005  Eudicot   Myrtaceae   myrtle   4,625  Eudicot   Apocynaceae   dogbane   4,555  Monocot   Cyperaceae   sedge   4,350  Eudicot   Malvaceae   mallow   4,225  Monocot   Araceae   arum   4,025  Eudicot   Ericaceae   heath   3,995  Eudicot   Gesneriaceae   gesneriad   3,870  Eudicot   Apiaceae or Umbelliferae   parsley   3,780  Eudicot   Brassicaceae or Cruciferae   cabbage   3,710 Magnoliid dicot   Piperaceae   pepper   3,600  Monocot   Bromeliaceae   bromeliad   3,540  Eudicot   Acanthaceae   acanthus   3,500  Eudicot   Rosaceae   rose   2,830  Eudicot   Boraginaceae   borage   2,740  Eudicot   Urticaceae   nettle   2,625  Eudicot   Ranunculaceae   buttercup   2,525  Magnoliid dicot   Lauraceae   laurel   2,500 

 Evolution 

 History of classification 

thumb|upright|From 1736, an illustration of Linnaean classification

The botanical term "angiosperm", from Greek words  ( 'bottle, vessel') and  ( 'seed'), was coined in the form "Angiospermae" by Paul Hermann in 1690, including only flowering plants whose seeds were enclosed in capsules. The term angiosperm fundamentally changed in meaning in 1827 with Robert Brown, when angiosperm came to mean a seed plant with enclosed ovules. In 1851, with Wilhelm Hofmeister's work on embryo-sacs, Angiosperm came to have its modern meaning of all the flowering plants including Dicotyledons and Monocotyledons. The APG system treats the flowering plants as an unranked clade without a formal Latin name (angiosperms). A formal classification was published alongside the 2009 revision in which the flowering plants rank as the subclass Magnoliidae. From 1998, the Angiosperm Phylogeny Group (APG) has reclassified the angiosperms, with updates in the APG II system in 2003, the APG III system in 2009, and the APG IV system in 2016.

 Phylogeny 

 External 

In 2019, a molecular phylogeny of plants placed the flowering plants in their evolutionary context:

 Internal 

The major groups of living angiosperms are:

Detailed Cladogram of the 2016 Angiosperm Phylogeny Group (APG) IV classification.

 Fossil history 

thumb|upright|Adaptive radiation in the Cretaceous created many flowering plants, such as Sagaria in the Ranunculaceae.

Fossilised spores suggest that land plants (embryophytes) have existed for at least 475 million years. However, angiosperms appear suddenly and in great diversity in the fossil record in the Early Cretaceous (~130 mya). Claimed records of flowering plants prior to this are not widely accepted. Molecular evidence suggests that the ancestors of angiosperms diverged from the gymnosperms during the late Devonian, about 365 million years ago. The origin time of the crown group of flowering plants remains contentious. By the Late Cretaceous, angiosperms appear to have dominated environments formerly occupied by ferns and gymnosperms. Large canopy-forming trees replaced conifers as the dominant trees close to the end of the Cretaceous, 66 million years ago. The radiation of herbaceous angiosperms occurred much later.

 Reproduction 

 Flowers 

thumb|upright=2|Angiosperm flower showing reproductive parts and life cycle

The characteristic feature of angiosperms is the flower. Its function is to ensure fertilization of the ovule and development of fruit containing seeds. It may arise terminally on a shoot or from the axil of a leaf. The flower-bearing part of the plant is usually sharply distinguished from the leaf-bearing part, and forms a branch-system called an inflorescence.

Flowers produce two kinds of reproductive cells. Microspores, which divide to become pollen grains, are the male cells; they are borne in the stamens. The female cells, megaspores, divide to become the egg cell. They are contained in the ovule and enclosed in the carpel; one or more carpels form the pistil.

The flower may consist only of these parts, as in wind-pollinated plants like the willow, where each flower comprises only a few stamens or two carpels. In insect- or bird-pollinated plants, other structures protect the sporophylls and attract pollinators. The individual members of these surrounding structures are known as sepals and petals (or tepals in flowers such as Magnolia where sepals and petals are not distinguishable from each other). The outer series (calyx of sepals) is usually green and leaf-like, and functions to protect the rest of the flower, especially the bud. The inner series (corolla of petals) is, in general, white or brightly colored, is more delicate in structure, and attracts pollinators by colour, scent, and nectar. 

Most flowers are hermaphrodite, producing both pollen and ovules in the same flower, but some use other devices to reduce self-fertilization. Heteromorphic flowers have carpels and stamens of differing lengths, so animal pollinators cannot easily transfer pollen between them. Homomorphic flowers may use a biochemical self-incompatibility to discriminate between self and non-self pollen grains. Dioecious plants such as holly have male and female flowers on separate plants. Monoecious plants have separate male and female flowers on the same plant; these are often wind-pollinated, as in maize, but include some insect-pollinated plants such as Cucurbita squashes.

 Fertilisation and embryogenesis 

Double fertilization requires two sperm cells to fertilise cells in the ovule. A pollen grain sticks to the stigma at the top of the pistil, germinates, and grows a long pollen tube. A haploid generative cell travels down the tube behind the tube nucleus. The generative cell divides by mitosis to produce two haploid (n) sperm cells. The pollen tube grows from the stigma, down the style and into the ovary. When it reaches the micropyle of the ovule, it digests its way into one of the synergids, releasing its contents including the sperm cells. The synergid that the cells were released into degenerates; one sperm makes its way to fertilise the egg cell, producing a diploid (2n) zygote. The second sperm cell fuses with both central cell nuclei, producing a triploid (3n) cell. The zygote develops into an embryo; the triploid cell develops into the endosperm, the embryo's food supply. The ovary develops into a fruit. and each ovule into a seed.

 Fruit and seed 

thumb|The fruit of the horse chestnut tree, showing the large seed inside the fruit, which is dehiscing or splitting open. 

As the embryo and endosperm develop, the wall of the embryo sac enlarges and combines with the nucellus and integument to form the seed coat. The ovary wall develops to form the fruit or pericarp, whose form is closely associated with type of seed dispersal system.

Other parts of the flower often contribute to forming the fruit. For example, in the apple, the hypanthium forms the edible flesh, surrounding the ovaries which form the tough cases around the seeds.

Apomixis, setting seed without fertilization, is found naturally in about 2.2% of angiosperm genera. Some angiosperms, including many citrus varieties, are able to produce fruits through a type of apomixis called nucellar embryony.

 Interactions with humans 

 Practical uses 

thumb|Harvesting rice in Arkansas, 2020

thumb|Food from plants: a dish of Dal tadka, Indian lentil soup

Agriculture is almost entirely dependent on angiosperms, which provide virtually all plant-based food and livestock feed. Much of this food derives from a small number of flowering plant families. For instance, half of the world's calorie intake is supplied by just three plants - wheat, rice and maize.

+ Major food-providing families Family  English  Example foods from that family Poaceae  Grasses, cereals  Most feedstocks, inc. rice, maize, wheat, barley, rye, oats, pearl millet, sugar cane, sorghum Fabaceae  Legumes, pea family  Peas, beans, lentils; for animal feed, clover, alfalfa Solanaceae  Nightshade family  Potatoes, tomatoes, peppers, aubergines Cucurbitaceae  Gourd family  Squashes, cucumbers, pumpkins, melons Brassicaceae  Cabbage family  Cabbage and its varieties, e.g. Brussels sprout, broccoli; mustard; oilseed rape Apiaceae  Parsley family  Parsnip, carrot, parsley, coriander, fennel, cumin, caraway Rutaceae  Rue family  Oranges, lemons, grapefruits Rosaceae  Rose family  Apples, pears, cherries, apricots, plums, peaches

Flowering plants provide a diverse range of materials in the form of wood, paper, fibers such as cotton, flax, and hemp, medicines such as digoxin and opioids, and decorative and landscaping plants. Coffee and hot chocolate are beverages from flowering plants.

 Cultural uses 

thumb|upright| Bird-and-flower painting: Kingfisher and iris kachō-e woodblock print by Ohara Koson (late 19th century) 

Both real and fictitious plants play a wide variety of roles in literature and film. Flowers are the subjects of many poems by poets such as William Blake, Robert Frost, and Rabindranath Tagore. Bird-and-flower painting () is a kind of Chinese painting that celebrates the beauty of flowering plants. Flowers have been used in literature to convey meaning by authors including William Shakespeare. 
Flowers are used in a variety of art forms which arrange cut or living plants, such as bonsai, ikebana, and flower arranging. Ornamental plants have sometimes changed the course of history, as in tulipomania. Many countries and regions have floral emblems; a survey of 70 of these found that the most popular flowering plant family for such emblems is Orchidaceae at 15.7% (11 emblems), followed by Fabaceae at 10% (7 emblems), and Asparagaceae, Asteraceae, and Rosaceae all at 5.7% (4 emblems each).

 Conservation 

thumb|left|upright=0.9|Viola calcarata, a species highly vulnerable to climate change.

Human impact on the environment has driven a range of species extinct and is threatening even more today. Multiple organizations such as IUCN and Royal Botanic Gardens, Kew suggest that around 40% of plant species are threatened with extinction. The majority are threatened by habitat loss, but activities such as logging of wild timber trees and collection of medicinal plants, or the introduction of non-native invasive species, also play a role. Relatively few plant diversity assessments currently consider climate change, yet it is starting to impact plants as well: according to IPCC, about 3% of flowering plants are very likely to be driven extinct within a century at , and 10% at .Parmesan, C., M.D. Morecroft, Y. Trisurat et al. (2022) Chapter 2: Terrestrial and Freshwater Ecosystems and Their Services in  In worst-case scenarios, half of all tree species, or of all plant species, may be driven extinct by climate change over that timeframe. 

Conservation in this context is the attempt to prevent extinction, whether in situ by protecting plants and their habitats in the wild, or ex situ in seed banks or as living plants. Some 3000 botanic gardens around the world maintain living plants, including over 40% of the species known to be threatened, as an "insurance policy against extinction in the wild." The United Nations' Global Strategy for Plant Conservation asserts that "without plants, there is no life". It aims to "halt the continuing loss of plant diversity" throughout the world.

 References 

Bibliography

Articles, books and chapters

 
 
 
 
 
 
 1st edition published by Oxford University Press in 1991 
 
 
 
 Cromie, William J. (December 16, 1999). "Oldest Known Flowering Plants Identified By Genes". Harvard University Gazette.
 
 
 
 
 
 
 
 

 
 
 
 
 
 
 
 

Websites

 
 
 

Angiosperms
Category:Plant sexuality
Category:Plants
Category:Pollination