Wikipedia

Poaceae

(Redirected fromGrass)
"Grass" redirects here. For other uses, seeGrass (disambiguation).
"Gramineae" redirects here. Not to be confused withGraminea.

Poaceae(/poʊˈeɪsi.iː,-ˌaɪ/), also calledGramineae(/ɡrəˈmɪni.iː,-ˌaɪ/), is a large and nearly ubiquitousfamilyofmonocotyledonousflowering plantscommonly known asgrasses.It includes thecerealgrasses,bamboos,the grasses of naturalgrasslandand species cultivated inlawnsandpasture.The latter are commonly referred to collectively asgrass.

Grasses
Temporal range:Albian–Present[1]
Flowering head of meadow foxtail (Alopecurus pratensis), with stamens exerted at anthesis
Scientific classificationEdit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Clade: Commelinids
Order: Poales
Clade: Graminid clade
Family: Poaceae
Barnhart[2]
Type genus
Poa
Subfamilies
Synonyms[3]

GramineaeJuss.

With around 780generaand around 12,000 species,[4]the Poaceae is the fifth-largestplant family,following theAsteraceae,Orchidaceae,FabaceaeandRubiaceae.[5]

The Poaceae are the most economically important plant family, providingstaple foodsfrom domesticatedcerealcrops such asmaize,wheat,rice,oats,barley,andmilletfor people and asfeedformeat-producing animals.They provide, through direct human consumption, just over one-half (51%) of all dietary energy; rice provides 20%,[6]wheat supplies 20%, maize (corn) 5.5%, and othergrains6%.[citation needed]Some members of the Poaceae are used as building materials (bamboo,thatch,andstraw); others can provide a source ofbiofuel,primarily via the conversion of maize toethanol.

Grasses havestemsthat are hollow except at thenodesand narrow alternate leaves borne in two ranks. The lower part of each leaf encloses the stem, forming a leaf-sheath. The leaf grows from the base of the blade, an adaptation allowing it to cope with frequent grazing.

Grasslandssuch assavannahandprairiewhere grasses are dominant are estimated to constitute 40.5% of the land area of theEarth,excludingGreenlandandAntarctica.[7]Grasses are also an important part of the vegetation in many other habitats, includingwetlands,forestsandtundra.

Though they are commonly called "grasses", groups such as theseagrasses,rushesandsedgesfall outside this family. The rushes and sedges are related to the Poaceae, being members of theorderPoales,but the seagrasses are members of the orderAlismatales.However, all of them belong to themonocotgroup of plants.

Description

Diagram of a typical lawn grass plant

Grasses may beannualorperennialherbs,[8]: 10 generally with the following characteristics (the image gallery can be used for reference): Thestemsof grasses, calledculms,are usually cylindrical (more rarely flattened, but not 3-angled) and are hollow, plugged at thenodes,where the leaves are attached.[8][9]Grassleavesare nearly always alternate and distichous (in one plane), and have parallel veins.[8]: 11 Each leaf is differentiated into a lower sheath hugging the stem and a blade with entire (i.e., smooth) margins.[8]: 11 The leaf blades of many grasses are hardened withsilicaphytoliths,which discourage grazing animals; some, such assword grass,are sharp enough to cut human skin. A membranous appendage or fringe of hairs called theligulelies at the junction between sheath and blade, preventing water or insects from penetrating into the sheath.[8]: 11 

Inflorecence scheme and floral diagram. 1 – glume, 2 – lemma, 3 – awn, 4 – palea, 5 – lodicules, 6 – stamens, 7 – ovary, 8 – styles.

Flowersof Poaceae are characteristically arranged inspikelets,each having one or more florets.[8]: 12 The spikelets are further grouped intopanicles or spikes.The part of the spikelet that bears the florets is called the rachilla. A spikelet consists of two (or sometimes fewer)bractsat the base, calledglumes,followed by one or more florets.[8]: 13 A floret consists of the flower surrounded by two bracts, one external—thelemma—and one internal—thepalea.The flowers are usuallyhermaphroditic—maizebeing an important exception—and mainlyanemophilousor wind-pollinated, although insects occasionally play a role.[10]Theperianthis reduced to two scales, calledlodicules,[8]: 11 that expand and contract to spread the lemma and palea; these are generally interpreted to be modified sepals. Thefruitof grasses is acaryopsis,in which the seed coat is fused to the fruit wall.[8]: 16  Atilleris a leafy shoot other than the first shoot produced from the seed.[8]: 11 

Growth and development

Grass flowers

Grass blades grow at the base of the blade and not from elongated stem tips. This low growth point evolved in response to grazing animals and allows grasses to begrazedormownregularly without severe damage to the plant.[11]: 113–114 

Three general classifications of growth habit present in grasses: bunch-type (also called caespitose),stoloniferous,andrhizomatous.[12] The success of the grasses lies in part in their morphology and growth processes and in part in their physiological diversity. There are bothC3andC4grasses, referring to the photosynthetic pathway for carbon fixation. The C4 grasses have a photosynthetic pathway, linked to specializedKranz leaf anatomy,which allows for increasedwater use efficiency,rendering them better adapted to hot, arid environments.[13]

The C3 grasses are referred to as "cool-season" grasses, while the C4 plants are considered "warm-season" grasses.[8]: 18–19 

Although the C4 species are all in thePACMAD clade(see diagram below), it seems that various forms of C4 have arisen some twenty or more times, in various subfamilies or genera. In theAristidagenus for example, one species (A. longifolia) is C3 but the approximately 300 other species are C4. As another example, the whole tribe ofAndropogoneae,which includesmaize,sorghum,sugar cane,"Job's tears",andbluestem grasses,is C4.[14]Around 46 percent of grass species are C4 plants.[15]

Taxonomy

The name Poaceae was given byJohn Hendley Barnhartin 1895,[16]: 7 based on the tribe Poeae described in 1814 byRobert Brown,and the type genusPoadescribed in 1753 byCarl Linnaeus.The term is derived from the Ancient Greekπόα (póa, "fodder" ).

Evolutionary history

Grasses include some of the most versatileplant life-forms.They became widespread toward the end of theCretaceousperiod, and fossilizeddinosaurdung (coprolites) have been found containingphytolithsof a variety that include grasses that are related to modernriceandbamboo.[17]Grasses have adapted to conditions in lushrain forests,drydeserts,cold mountains and evenintertidal habitats,and are currently the most widespread plant type; grass is a valuable source of food and energy for all sorts of wildlife.

A cladogram shows subfamilies and approximate species numbers in brackets:[14]

Drawing ofAnomochloa marantoidea,one of the most primitive living grass species

Before 2005, fossil findings indicated that grasses evolved around 55 million years ago. Finds of grass-likephytolithsinCretaceousdinosaurcoprolitesfrom the latest Cretaceous (Maastrichtian) agedLameta Formationof India have pushed this date back to 66 million years ago.[18][19]In 2011, fossils from the same deposit were found to belong to the modern rice tribeOryzeae,suggesting substantial diversification of major lineages by this time.[20]

In 2018, a study described grass microfossils extracted from the teeth of thehadrosauroiddinosaurEquijubusnormanifrom northern China, dating to theAlbianstage of theEarly Cretaceousapproximately 113–100 million years ago, which were found to belong to primitive lineages within Poaceae, similar in position to the Anomochlooideae. These are currently the oldest known grass fossils.[1]

The relationships among the three subfamilies Bambusoideae, Oryzoideae and Pooideae in the BOP clade have been resolved: Bambusoideae and Pooideae are more closely related to each other than to Oryzoideae.[21]This separation occurred within the relatively short time span of about 4 million years.

According toLester Charles King,the spread of grasses in theLate Cenozoicwould have changed patterns ofhillslope evolutionfavouring slopes that are convex upslope and concave downslope and lacking afree facewere common. King argued that this was the result of more slowly acting surface wash caused by carpets of grass which in turn would have resulted in relatively moresoil creep.[22][23]

Subdivisions

There are about 12,000 grass species in about 771 genera that are classified into 12 subfamilies.[24]See the fulllist of Poaceae genera.

Setaria verticillatafromPanicoideae

Distribution

The grass family is one of the most widely distributed and abundant groups of plants onEarth.Grasses are found on every continent,[25][26]includingAntarctica.The Antarctic hair grass,Deschampsia antarcticais one of only two plant species native to the westernAntarctic Peninsula.

Ecology

Wind-blown grass in theValles Calderain New Mexico, United States

Grasses are thedominantvegetation in many habitats, includinggrassland,salt-marsh,reedswampandsteppes.They also occur as a smaller part of the vegetation in almost every other terrestrial habitat.[citation needed] Grass-dominatedbiomesare called grasslands. If only large, contiguous areas of grasslands are counted, these biomes cover 31% of the planet's land.[27]Grasslands includepampas,steppes,andprairies.[28] Grasses provide food to manygrazingmammals,[29]as well as to many species ofbutterfliesandmoths.[30][31] Many types of animals eat grass as their main source of food, and are calledgraminivores– these includecattle,sheep,horses,rabbitsand manyinvertebrates,such asgrasshoppersand the caterpillars of manybrown butterflies.Grasses are also eaten byomnivorousor even occasionally by primarilycarnivorousanimals.

Akangarooeating grass

Grasses dominate certainbiomes,especiallytemperate grasslands,because many species are adapted to grazing and fire.[32]

Grasses are unusual in that themeristemis near the bottom of the plant; hence, grasses can quickly recover from cropping at the top.[33] The evolution of large grazing animals in theCenozoiccontributed to the spread of grasses. Without large grazers, fire-cleared areas are quickly colonized by grasses, and with enough rain, tree seedlings. Trees eventually outcompete most grasses. Trampling grazers kill seedling trees but not grasses.[11]: 137 

Sexual reproduction and meiosis

Sexual reproductionandmeiosishave been studied inrice,maize,wheatandbarley.[34]Meiosis research in these crop species is linked to crop improvement, since meioticrecombinationis an important component ofplant breeding.[34]Unlike in animals, the specification of both male and female plantgermlinesoccurs late in development during flowering. The transition from thesporophytephase to thegametophytestate is initiated by meiotic entry.[34]

Uses

Grasses are, in human terms, perhaps the most economically important plant family.[35]Their economic importance stems from several areas, including food production, industry, andlawns.They have been grown as food fordomesticated animalsfor up to 6,000 years[citation needed]and the grains of grasses such aswheat,rice, maize (corn) andbarleyhave been the most important humanfood crops.Grasses are also used in the manufacture ofthatch,paper,fuel,clothing,insulation,timber forfencing,furniture,scaffoldingandconstructionmaterials, floormatting,sports turfandbaskets.

Grazing cattle on a pasture nearHradec nad MoravicíinCzech Silesia.

Food production

Of all crops grown, 70% are grasses.[36]Agricultural grasses grown for their edible seeds are calledcerealsorgrains(although the latter term, when used agriculturally, refers to both cereals and similar seeds of other plant species, such asbuckwheatandlegumes). Three cereals—rice, wheat, and maize (corn)—provide more than half of all calories consumed by humans.[37]Cereals constitute the major source ofcarbohydratesfor humans and perhaps the major source of protein; these include rice (insouthernandeastern Asia), maize (inCentralandSouth America), and wheat and barley (inEurope,northern Asiaand theAmericas).

Sugarcaneis the major source ofsugarproduction. Additional food uses of sugarcane includesprouted grain,shoots,andrhizomes,and in drink they includesugarcane juiceandplant milk,as well asrum,beer,whisky,andvodka.

Bamboo shootsare used in numerous Asian dishes and broths, and are available in supermarkets in various sliced forms, in both fresh, fermented and canned versions.

Lemongrassis a grass used as a culinary herb for its citrus-like flavor and scent.

Many species of grass are grown aspasturefor foraging or asfodderfor prescribedlivestockfeeds, particularly in the case ofcattle,horses,andsheep.Such grasses may be cut and stored for later feeding, especially for the winter, in the form of bales ofhayorstraw,or in silos assilage.Straw (and sometimes hay) may also be used as bedding for animals.

An example of asod-forming perennial grass used in agriculture isThinopyrum intermedium.

Industry

Grasses are used as raw material for a multitude of purposes, including construction and in the composition of building materials such ascob,for insulation, in the manufacture of paper and board such asoriented structural straw board.Grassfibercan be used for makingpaper,biofuelproduction,[38]nonwoven fabrics, and as replacement for glass fibers used in reinforced plastics.[39]Bamboo scaffoldingis able to withstand typhoon-force winds that would break steel scaffolding.[27]Larger bamboos andArundo donaxhave stout culms that can be used in a manner similar to timber,Arundois used to make reeds forwoodwind instruments,and bamboo is used for innumerable implements.[40]

Phragmites australis(common reed) is important forthatchingand wall construction of homes in Africa.[41]Grasses are used inwater treatmentsystems,[42]inwetland conservationandland reclamation,and used to lessen the erosional impact of urban storm water runoff.[43]

Palaeoecological reconstructions

Processed, fossilised pollen from the family Poaceae. Species unknown

Pollenmorphology, particularly in thePoaceaefamily, is key to figuring out their evolutionary relationships andhow environments have changed over time.[44]Grass pollen grains, however, often look the same, making it hard to use them for detailed climate or environmental reconstructions.[45][46]Grass pollen has asingle poreand can vary a lot in size, from about 20 to over 100 micrometers, and this size difference has been looked into for clues about past habitats, to tell apartdomesticatedgrasses from wild ones,[47][48][49]and to indicate various biological features like how they performphotosynthesis,[50]their breeding systems,[50][51]and genetic complexity.[52][53][45]Yet, there's ongoing debate about how effective pollen size is for piecing together historical landscapes and weather patterns, considering other factors such as genetic material amount might also affect pollen size.[54][55][45]Despite these challenges, new techniques inFourier-Transform Infrared Spectroscopy(FT-IR) and improved statistical methods are now helping to better identify these similar-looking pollen types.[46]

Lawn and ornamental use

Main articles:LawnandOrnamental grass
A lawn in front of a building

Grasses are the primary plants used in lawns, which themselves derive from grazedgrasslandsin Europe.[citation needed]They also provide an important means of erosion control (e.g., along roadsides), especially on sloping land.[citation needed]Grass lawns are an important covering of playing surfaces in many sports, includingfootball (soccer),American football,tennis,golf,cricket,softballandbaseball.

Ornamental grasses, such asperennialbunch grasses,are used in many styles ofgarden designfor their foliage, inflorescences and seed heads. They are often used innatural landscaping,xeriscapingand slope and beach stabilization in contemporary landscaping,wildlife gardening,andnative plant gardening.[citation needed]They are used as screens and hedges.[56]

Sports turf

See also:Turf managementandSand-based athletic fields

Grass playing fields, courses and pitches are the traditional playing surfaces for manysports,includingAmerican football,association football,baseball,cricket,golf,andrugby.Grass surfaces are also sometimes used forhorse racingandtennis.Type of maintenance and species of grass used may be important factors for some sports, less critical for others. In some sports facilities, including indoor domes and other places where maintenance of a grass field would be difficult, grass may be replaced withartificial turf,a synthetic grass-like substitute.[57]

Cricket

The gray area is thecricketpitch currently in use. Parallel to it are other pitches in various states of preparation which could be used in other matches.
Main article:Cricket pitch § Preparation and maintenance of the playing area

In cricket, the pitch is the strip of carefully mowed and rolled grass where the bowler bowls. In the days leading up to the match it is repeatedly mowed and rolled to produce a very hard, flat surface for the ball to bounce off.[58]

Golf

Main article:Golf course

Grass on golf courses is kept in three distinct conditions: that of therough,thefairway,and theputting green.Grass on the fairway is mown short and even, allowing the player to strike the ball cleanly. Playing from the rough is a disadvantage because the long grass may affect the flight of the ball. Grass on the putting green is the shortest and most even, ideally allowing the ball to roll smoothly over the surface. An entire industry revolves around the development and marketing of turf grass varieties.[59]

Tennis

Main article:Grass court

In tennis, grass is grown on very hard-packed soil, and the bounce of atennis ballmay vary depending on the grass's health, how recently it has been mowed, and the wear and tear of recent play.[citation needed]The surface is softer thanhard courtsandclay(other tennis surfaces), so the ball bounces lower, and players must reach the ball faster resulting in a different style of play which may suit some players more than others.[citation needed]Among the world's most prestigious court for grass tennis is Centre Court atWimbledon, Londonwhich hosts the final of the annualWimbledon Championshipsin England, one of the fourGrand Slamtournaments.

Economically important grasses

Grain crops
Leaf and stem crops
Lawn grasses
Ornamental grasses(Horticultural)
Model organisms

A number of grasses are invasive species that damage natural ecosystems, including forms ofPhragmites australiswhich are native to Eurasia but has spread around the world.[60][61]


Role in society

Grass-covered house inIceland
Typical grass seen in meadows
Grass with non-grass flowers around it

Grasses have long had significance in human society. They have been cultivated as feed for people anddomesticated animalsfor thousands of years. The primary ingredient ofbeeris usually barley or wheat, both of which have been used for this purpose for over 4,000 years.[62]

In some places, particularly insuburbanareas, the maintenance of a grass lawn is a sign of a homeowner's responsibility to the overall appearance of their neighborhood. One work credits lawn maintenance to:

...the desire for upward mobility and its manifestation in the lawn. As Virginia Jenkins, author ofThe Lawn,put it quite bluntly, "Upper middle-class Americans emulated aristocratic society with their own small, semi-rural estates." In general, the lawn was one of the primary selling points of these new suburban homes, as it shifted social class designations from the equity and ubiquity of urban homes connected to the streets with the upper-middle class designation of a "healthy" green space and the status symbol that is the front lawn.[63][64]

In communities withdroughtproblems, watering of lawns may berestrictedto certain times of day or days of the week.[65]Many US municipalities and homeowners' associations have rules which require lawns to be maintained to certain specifications, sanctioning those who allow the grass to grow too long.[66][67]

Thesmell of freshly cut grassis produced mainly bycis-3-Hexenal.[68]

Some commonaphorismsinvolve grass. For example:

  • "The grass is always greener on the other side" suggests an alternate state of affairs will always seem preferable to one's own.
  • "Don't let the grass grow under your feet" tells someone to get moving.
  • "Asnakein the grass "means dangers that are hidden.
  • "Whenelephantsfight, it is the grass which suffers "tells of bystanders caught in the crossfire.

A folk myth about grass is that it refuses to grow where any violent death has occurred.[69]

See also

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