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ANGIOSPERMS Angiosperms •Flowering plants,producingseedsthatarefullyenclosedbyfruits. •Theyarethemosthighlyevolvedofplants,andappearfirstinrocksof LowerCretaceousage(about130millionsyearsago). •AllofthemareincludedinthedivisionofAnthophyta (Magnoliophyta), comprisingabout250000species,fromverysmall(e.g.Lemna)tohuge ones(e.g.Eucalyptus). •Antophytaisdividedintwomainclasses:Monocotyledones (Liliopsida) withabout65.000speciesandDicotyledones (Magnoliopsida)withabout 165.000species. •Theirgreatimportancederivesfromthequantity,varietyandactivityof activecompounds(secondarymetabolites)thatisthereasonofthe phytopharmaceuticalstudy.Themajorityofmedicinalplantsare angiospermdicotyledones. Angiosperms •Theyaretheplantscapabletoadapttoall theenvironmentsandtowinthecompetition withotherplantorganismsthanksto: vflower vfruit Magnoliophyta Eucalyptus Lemna Evolutionarytrendsfromgymnospermsto angiosperms •Slow-growingtreespecies(gymnosperms)are displacedbyfast-growingherbs(mostevolved angiosperms)capabletoincreaserapidlytheir populations. •Thelossof‘woodiness’,infavorof‘herb-ness’, correspondstotheabandonofthemechanical defenses againstherbivoresandpathogensin favorofmoreselectiveandeffectivechemical defenses,basedontheproductionofactive compounds. Trachea,typicalof angiosperms Tracheid,typical ofgymnosperms Fromgymnospermstoangiosperms •Improvement of vascular tissues (appearing of tracheae or vessels) played an important role in the diffusion of angiosperms since it allowed an improvement of photosynthetic activity, a higher production of organic compounds and a faster growth. Whyangiospermsarethemost evolvedplants? ØHigh rate of pollination and cross-fertilization assured by insects and animal pollinators that are attracted to flowers. ØPresence of a food-storing endosperm in the seed allowing to survive under adverse environmental conditions. ØHigh rate of seed dispersion assured by the fruit. ØEfficient vascular system owing to the presence of tracheae into xylem and of sieve tubes into phloem. ØLeaf apparatus more developed and efficient for photosynthesis. ØDeciduous habitus allows to survive during adverse conditions. ØProduction of a huge amount of secondary metabolites efficient in the defense and propagation of the plant species. MonocotyledonesvsDicotyledones Liliopsida atactostele Magnoliopsida eustele Tetrarchrootofa dycotyledone Polyarchrootofa monocotyledone dicotyledones monocotyledones Stomatadistribution Cuscuta Monotropa •Almostallangiospermsareautotrophous,butthereareafewparasites (livingonotherplantstowhichtheycancauseseriousdamage)or saprophytes(absorbingsolubleorganicnutrientsfromdeadplant,animal matter,dung,etc.). Asteridae Rosidae Dicotyledones Magnolidae Monocotyledones Protoangiospermae Classificationaccording toCronquist(1981) Gnetales Gimnospermae Magnoliopsida(dicotyledones) • • • • Seedwith2embryoleaves(orcotyledons). Woodyorherbaceousplants. Presenceofataproot. Eustelicprimarythickeningofstem,withopened, collateralbundles. • Heteroxilwoodwithtracheidsandtracheae. • Flowerwithcalyxandcorollaandwhorlsof4-5elements. Evolutionarytrendsinangiosperms •Indicotyledones thereisanabandonofslow-growingandelderly treespecies(derivedfromGymnosperms)infavouroffast-growing andshort-livingherbs,capableofproducingagreatnumberof individualsinapopulations. •Monocotyledonegroupsseemtobeindipendenteachother(e.g. Orchidaceaewithcomplexandshowyflowers,Graminaewith modestflowers,Palmaewitharboreoushabitus,Liliaceaewitha greatvarietyofherbaceousformsetc.). Magnolidae • TransitiongroupfromGymnospermsatthestartof Angiospermsevolution. • Woodyplantswidespreadintheoldestforests(e.g. Amazon). • Initialprevalenceoftracheids. • Flowerswithoutpetalsorwithmanypetalsusually spiral-inserted. • Atphytochemicallevel,theyproduceaminoacids derivativessuchasalkaloidsandphenoliccompounds derivedfromshikimicacid(e.g.simplephenols, gallotannins). • Theyincludeabout20familiesamongwhich Magnoliaceae,Lauraceae,Piperaceae, Aristolochiaceae,Nympaeaceae etc. Rosidae • Theyabandonthewoodyhabitusinfavourof shrub/herbaceoushabitus. • Flowermorecomplex:2groupsofspecializedfloral leaves(regular,actinomorphicflowersradially symmetrical;calyxwithsepalsandcorollawith5 separate,showyandmorphologicallysimilarpetals). • Zygomorphicflowers(i.e.bilaterallysymmetricaland thereforedivisibleintoequalhalvesinonly1plane)in themostevolutedforms. • Phenylpropanidunits,previouslyusedforthesynthesis oflignins,arenowusedtoproducecoumarins,tannins andflavonoids.Atthebeginning,theproductionof terpenoids,whilelittlethatofalkaloids. Asteridae • Maximumistheherbaceouscharacterandshortthelife cycle. • Flowershapereachesthegreatestcomplexityasa consequenceofthemaximumspecializationtowards pollinators.Zygomorphicflowerwithtube-shapedcorolla, orarrangementofmoreflowersinparticularinflorescences. • Becauseoftheirwidesecondarymetabolism,theyhave colonizedmostoftheplanetandconstitutedanimportant sourceofactivecompoundstobeusedintherapy.The secondarymetabolitesarerepresentedmainlyby terpenoidsthatinthearomaticplantsreachtheirmaximum expression. Liliopsida(monocotyledones) • Seedwithonecotyledon. • Habitusonlyherbaceous.Lackingofthesecondary thickening(exceptPalmae). • Fibrousroots;presenceofadventitiousrootsonthestem. • Atactostelestructureofstem,withclosedvascularbundles randomlydistributed. • Simple,sessileleaves,withamplexicaulsheathandparallel nervation. • Flowerscomposedof3elements;nodistinctionbetween calyxandcorolla:sepalsandpetalsaredisplacedbytepals. • Unlikedicotyledones,thevariousgroupsaretaxonomically indipendent. • Secondarymetabolitesvariedbetweenthevariousplant groups.Anyway,theyarenotsonumerousasinthemost evolvedformsofdicotyledones. Angiospermsaretherichest sourceofactivecompounds •Thenumberofpharmacologically-activecompounds (drugs)increasesfromwaterorganisms(e.g.Algae)to terrestrialplants,aswellasfromphotosynthetically inefficientBryiophytestothoseefficient(e.g.broad-leaf Angiosperms). •Asamatteroffact,thehigherenergyderivedfrom photosynthesisandthebetterphysiologicalefficiency, stimulatedthesyntheticpotentialofplantssothatthey interactwiththeenvironmentthroughtheproductionofa greatnumberofsecondarymetabolites. ClassificationaccordingtoCronquist(1981) Magnolidae •Monimiaceae •Lauraceae •Illiciacee •Papaveraceae Rosidae •Rosaceae •Mimosaceae •Fabaceae •Caesalpiniaceae •Myrtaceae •Euphorbiaceae •Rhamnaceae •Hippocastanaceae •Rutaceae •Araliaceae Monocotyledones(Liliopsida) •Apiaceae •Liliaceae •Poaceae •Zingiberaceae •Iridaceae •Aloeaceae Asteridae •Gentianaceae •Apocynaceae •Solanaceae •Lamiaceae •Plantaginaceae •Scrophulariaceae •Rubiaceae •Valerianaceae •Asteraceae Flower •Floweristhestructureconcernedwithsexualreproduction, consistingoftheandroecium (maleorgans:stamens)and gynoecium (femaleorgans:carpels),commonlysurroundedby sterilecorolla (petals)andcalyx (sepals). Flowerandreproduction •Floweriscomposedof4groupsofmodifiedleaves:sepals,petals,stamens andcarpels. •Sepals aretheouterfloralleaves,usuallygreenish,whichareborneinatight spiralorwhorled.Theyformcalyx whichprotectsflower(formingbuds)before opening.Sepalscanbeseparateorfused.Sometimescalyxispersistentand accompaniesfruitafterfertilization. •Petals aretheinnerfloralleaves,usuallybrightlycolouredorsmellingto attractpollinators,andborneinatightspiral,orwhorled.Theyformcorolla whichenclosesthereproductiveorgansand,togethercalyx,protectsthem. Petalscanbeseparateorfused.Afterfertilizationpetalsfalldown. •Stamens arefertileleaves(sporophylls)formingthemaleorganscomprising astalk(thefilament)andtheanthers whicharecommonly2-lobed.Anthers containmicrosporangia thatproducemicrospores givingrisetomale gametophytes. •Carpels arefertileleaves(sporophylls)formingthefemaleorganscomprising anovary (containing1tomanyovules borneonaplacenta)andwithausually terminalstyle tippedbythestigma.Ovulescontainmegasporangia that producemegaspores givingrisetofemale gametophytes(embryosac). Structureofflowers PERFECTorMONOCLINOUS (BISEXUAL) IMPERFECTorDICLINOUS (UNISEXUAL) Stamensandcarpelsoccurin thesameflower Stamensorcarpelsinseparate flowersinthesameplant MONOICIOUSPLANT DIOICIOUSPLANT Plant with bisexual flowers, or alternatively with both stamen and carpels in flowers in the same individuals Flowers with stamens and carpels in separate individuals COMPLETE INCOMPLETE Withsepals,petals,stamensand carpelsinthesameflower Lackofoneormorefloralparts Floralsymmetry Regularflowers (Actinomorphic) Irregularflowers (Zygomorphic) Similar elements ofwhorls,with radial symmetry,i.e.divisible into equal halves inmany planes Differentelementofwhorls,with bilateralsymmetry,i.e.divisibleinto equalhalvesinonly1plane Morphologyof calyxandcorolla Morphologyof calyxandcorolla Originofthecarpelfromareproductiveleaf (sporophyll) •Aleafbearingmegasporangiafoldedinordertoprotectthem. Inflorescences Monopodialorindefinite (lateralfloralbranchesarisefrom amain,centralaxis) •Racemeorcluster •Spike(spadix,catkin) •Corymb •Umbel •Capitulum Sympodialordefiniteor cymose (apparentmainaxismadeupof manylateralfloralbranches, eacharisingfromtheonebefore) •Monochasium •Dichasium Compound(mainaxiswithlateralbranchesbearing thesameoradifferenttypeofinflorescence) •Panicleorcompoundraceme •Compoundumbel •Polychasium •Spikeletorcompoundspike •Clusterofcapitula •Clusterofspikes Inflorescences panicle composite umbel spike raceme corymb helicoid monochasium dichasium scorpioid monochasium spadix umbel capitulum polychasium •Monopodial orindefinite inflorescences •Compound anddefiniteinflorescences Monopodial inflorescences •Raceme:themainaxiscontinuestogrow,producingflowerslaterally withpedicels. •Spike:aracemeinwhichtheflowersareallsessile;ischaracteristicof Poaceae. •Corymb:aracemoseinflorescenceinwhichthelowerpedicelsare longerthantheuppersothattheflowerslieasadome,ordish,and theoutlineisroundishorflattish. •Umbel:allthepedicelsariseattheapexofanaxis.Itiscommonly compound,usuallyumbrella-shaped,andischaracteristicofApiaceae. •Capitulum:consistsofcloselypackedflowersorfloretswhichhaveno stalksandariseonaflattenedaxis,allatthesamelevel.Itis surroundedbyaninvolucreofbractsgivingittheappearanceofa singleflower.CapitulaaretypicalofAsteraceae. Cymose andcompound inflorescences •Monochasium:consists ofasinglefloral branch bearing flowers andending inasingleterminalflower (e.g.scorpioid andhelicoid cymes). •Dichasium:each branch give risetotwo morebranches. •Polychasium :each branch give risetomany morebranches. •Panicle:acompoundraceme,i.e.one that comprises several racemoseparts; it is characteristic ofPoaceae. •Compoundumbel:anumbel inwhich thepedicels give risetoother umbels; it is typical ofApiaceae. •Spikelet orcompoundspike:aspike inwhich shortbranches give riseto2-3 spikes;it is characteristic ofPoaceae. •Clusterofcapitula:aracemoseinflorescence inwhich thepedicels give rise tomany capitula. •Clusterofspikes:aracemoseinflorescence inwhich thepedicels give riseto many spikes. Inflorescences Inflorescences Inflorescences •Capitulum:consistsofcloselypacked flowersorflorets(discandrayflorets) whichhavenostalksandariseona flattenedaxis,allatthesamelevel.Itis surroundedbyaninvolucreofbracts givingittheappearanceofasingleflower. CapitulaaretypicalofAsteraceae. Inflorescences Capitulum of thistle with only tubular, disc florets Capitulum of dandelion with only ligulate, ray florets Capitulum of sunflower with both tubular and ligulate florets Inflorescences •Compound umbel:all thepedicels arise attheapex ofan axis;each pedicel gives riseto one umbel.It is characteristic of Apiaceae. Inflorescences simpleumbel compoundumbel Inflorescences •SpikeletistypicalofPoaceae. Inflorescences •PaniclesofZeamays Development of gametophytes sporogenesis gametogenesis Stamens •Stamenisthemaleorganofa flower,comprisingastalk(the filament)andtheanther whichis commonly2-lobed,eachlobe containing2pollensacs (microsporangia) inwhich microsporocytes undergo2meiotic divisionstoproduce4microspores thatbecomepollengrains. immaturepollengrain Pollengrain •Pollengrainderivesfromgerminatingmicrospore.Itcontainsthe malegametophyte,astructuremadeupofthepollengrainplusa pollentube.Thegraincontains3haploid nuclei(atubenucleusand2 spermnuclei). Pollengrain • Pollengrain(20-250μm)isprotectedby anouterenvelopecalledexin (composed ofsporopollenin)showingvarioustypes ofsculptures(taxonomicvalue),andbyan innerenvelopecalledintin (composedof celluloseandpectins).Itbearsstretched orcircularopeningsfromwhichthepollen tubecomesout.Theyareknownassolcus andporus. • Pollengrainsareanimportantsourceof nutrientsforanimalsandinsects(richof starchandlipids). • Theyarefrequentinfossils. VariouskindsofpollengrainsatSEM Carpel •Carpel isthefemalereproductiveorgan oftheflower,i.e.aunitofthegynoecium, comprisinganovary (containing1tomany ovules borneonaplacenta)andwitha usuallyterminalstyle tippedbythe stigma. •Styleisanextensionofthecarpelwhich supports thestigma. •Stigmaisthepartonwhichpollengrains germinate. c a r p e l Pollinationandpollen tubeformation •Stigma isthepartonwhich pollengrainsgerminatethrough absorptionofwaterand consequentformationofthe pollentube. •Angiospermshaveastigmatic pollination. Pollinationandpollen tubeformation •Thepollen grain contains 3haploid nuclei (atubenucleus and2 sperm nuclei),which passdownthetube to theovum.One of thesperm nucleifertilizes theovum,and thesecond fuses with the2polar nuclei forming theendosperm.Thetubenucleus (which is considered to be vestigial,having been completely functional earlier inthe evolution offlowering plants)degenerates after double fertilization (socalled because ofthetwo unions ofnuclei). •With the appearance of flower, angiosperms have solved the problem related to the impossibility to move towards a reproductive partner. •Flower assures a high rate of cross pollination. •Pollination is thetransferofpollen grains (containing male gametophytes)fromtheanther tothestigmaofaflowering plant.Flowers andinsects have co-evolved. •Inabattle ofthree armies,Large Bluesbutterflies arelured to oregano plants where theinsect flyers lay their eggs.After two weeks,theconsequent caterpillarscrwal tothebottom oftheplant where they exude a scent that tricks Myrmica ants into carrying thebutterfly larvae into theundergroundcolony.The caterpillarsspend thenext 11 months feeding ontheant grubs andgaining 98%oftheir body mass,before ultimately killing the colony andescaping. •Rosemary’santhersdepositingwhitepollen grainsonhairybackofabee. •Flowerspollinatedbybeesareyelloworbluesince theseinsectsarenotsensitivetoredwavelenghts. •FlowerofDigitalis withtubular corollashowinginsidesomestains actingaslandingplatformforbees. •FlowersofthegenusOphrys (Orchidaceae),mimingtheshape ofafemalebee,attractmale pollinators. •Nectarsareanimportantfeedingsourcefor pollinators.Theyaresecretorystructures producingenergeticnutrients. •Flowerspollinatedbybirdsareoftenofredcolour. •Graminaehaveananemophilous pollination:hairystigmasgetpollen grainsfromhanginganthers. •Wind-pollinatedplantsdonot dependoninsectssothattheydonot spendenergyfornectarproduction, andarecolourlessandodourless. •Anemophilouspollinationisless efficientandsucceedsonlywhena highnumberofindividualsofa populationstaycloseeachother(no fartherthan100m). •Someaquaticangiospermsarepollinatedbywater. •Themostimportantpigmentsforthecolour offlowersareflavonoids (mainly anthocyanins)andcarotenoids. •Fertilizationinvolvesthefusionoftwohaploidnucleifromtwo distinctindividuals(cross-fertilization)orfromoneindividual(selffertilization). •Normallymaleandfemaleorgansofaflowermatureatdifferent times(dichogamy),oralternativelytheyarephysicallyseparatedin ordertofavourcross-fertilization. •Plantswithself-pollination havesmall,colourlessflowers sincetheydonotneedtoattractpollinators. •Self-pollinatedpopulationshavealowlevelofgenetic variability. •Self-pollinationisfavouredinsomecases(e.g.disturbed areaswherepollinatorsarerare). Femalegametophyteanddoublefertilization •Thefemalegametophyte,knownasembryosac,is anovalstructureinthenucellus oftheovule of floweringplants,formedbythedivisionofthe haploidmegaspore nucleus(whichisproducedby theonlydiploidcellundergoingmeiosisinthe megasporangium),andthesiteoffertilization ofthe egganddevelopmentoftheembryo. •Itconsistsof6haploidcellswithoutcellwalls(2 synergidae,3antipodalcells,andaneggcell)and2 haploidnuclei(polarnuclei).Sometimesthe2 haploid,polarnucleifusetoformasingle, endospermmothercell.Atfertilization,1male nucleusfuseswiththeeggnucleustoformazygote whichdevelopsintotheembryo.The secondmale nucleusfuseswiththeprimaryendospermnucleus toformtheendospermnucleus.Thisthendividesto formtheendosperm. Doublefertilization •It occurs only in angiosperms and is the production of 2 sperm nuclei, both of which contribute to fertilization. •The male gametophyte, comprising the pollen grain and pollen tube, contains 2 sperm nuclei and 1 vegetative nucleus. •The vegetative nucleus degenerates once the pollen tube has penetrated the embryo sac by micropyle. •The 2 sperm nuclei enter the embryo sac. One unites with the egg nucleus to form the zygote, which develops into the embryo. The other unites with both of the polar nuclei or with the nucleus formed by their fusion, to form the primary endosperm nucleus, from which endosperm develops. •Endosperm is a triploid tissue storing food materials which are broken down during seed germination. •Double fertilization assures endosperm development just in the case of egg cell fertilization, avoiding the waste of nutritive substances. Doublefertilization Seed •Seed ofangiospermsiscomposedof2 diploid generations(theouter integuments andtheembryo)andone triploid generation(endosperm). •Embryohasusually1or2cotyledons. •Endosperm functionsasstorageof nutrients(carbohydrates,proteins,lipids) forembryoandseedgermination. •Seednutrientsarestoredinthevacuole orleucoplasts ofendospermcells. Lifecycleofanangiosperm Fruit • After fertilization, ovule is transformed into seed, ovary into fruit. • Fruits protect quiescent seeds and contribute to their dispersal. Fruit • Ovarywallsknownaspericarp, thickenanddifferentiateinto3 layers:theinnerendocarp,the middlemesocarp andtheouter exocarp.Theselayersareusually moreevidentinfleshyfruits. Classificationoffruits origin AGGREGATE SIMPLE MULTIPLE formed by the joining of several carpels that were separate in the flower (e.g. Rubus). formed from a single carpel or from various fused carpels formed from inflorescence pineapple) dehiscence INDEHISCENTremainingclosedat maturity openingnaturallyatmaturity follicle,legume,lomentum,siliqua, capsule,schizocarp consistence FLESHY(oneor DRY(oneseed) berry,drupe,pome samara,nut,achene, caryopsis moreseeds) DEHISCENT(many-seeded,dry) an entire (e.g. fig, Classificationoffruits TRUE Strictly,theripened ovaryofaplantandits contents FALSE Whentheripenedovary andseedsarecombined withotherstructures;e.g. theapple(apome)inwhich thetruefruit(core)is surroundedbyfleshderived fromthefloralreceptacle. Fleshyfruits • Drupe: a fleshy indehiscent fruit containing one or a few seeds, each enclosed within a stony endocarp, as in the genus Prunus (Rosaceae). Fleshyfruits • Berry:afleshy,indehiscent,fruitcontainingmanyseeds immersedinapulp(endocarp+mesocarp);theydonothave hardpartsexcepttheseeds(e.g.grape,tomatoes,blueberry, currant,banana,avocado,pepper,eggplant,belladonna). Fleshyfruits • Hesperidium:aberryinwhichthefleshypartisdividedinto segmentsandtheouterskinisatough,glandularleathery rind,e.g.Citrus(Rutaceaesuchasorange,lemonetc.). Fleshyfruits • Pepper:aberry,typicalofCapsicum wherethefleshy pericarpsurroundsahollowcontainingmanyseeds. Fleshyfruits • Pepo:aunilocular,many-seeded,hard-walledberrythat formsthefruitofCucurbitaceae(e.g.marrow,melon, watermelon,cucumberetc.). Fleshyfruits • Sorosis:afleshy,aggregatefruit(e.g.mulberry,currantetc.); strawberryisanaggregate,falsefruit,wherethepulp originatesfromreceptaclewhilethetruefruitsareachenes. Fleshyfruits (enlargementof thereceptacle) • Pome:fruitinwhichtheseedsareprotectedbyatoughcarpelwall andtheentirefruitisembeddedinafleshyreceptacle;inthe apple,thecarpelwallsurroundingtheseedscomprisesthecore, whichisthefruit,theediblefleshypartoftheapplebeingthe receptacle. Fleshyfruits • Syncarp:afleshy,multiplefruitformedfroman inflorescence,oftenincludingbracts,asinpineapple. Ripening immature • Ripeningisastageinthe developmentoffruit,characterized bythesoftening,colouring,and sweeteningofthetissue,andbya decreaseinacidity.Oftenthisis associatedwithanincreaseinthe ratesofrespiration,cellexpansion, andethylene production,combined withalossofchlorophyll. • Thecolourchangeisasignalforthe seeddispersors. ripening Ripening • Redcolourisdominantamongripefruitssince animals,thataresensitivetoredwavelenghts,are thebestseeddispersors. Dryfruits • Follicle:adryfruit derived from asingle carpel,containing morethan one seed, which dehisces along one side (along the suture)only. • Siliqua:thecapsularfruit ofthe Brassicaceae when atleast three times as longas broad,containing many seeds attached to acentral outgrowth known as replum;it dehisces along dorsal andventral sides (sutures). Dehiscentfruits • Capsule:adry,dehyscentfruit formedfromasyncarpous ovary;itcontainsmanyseeds andopensindifferentways. Dehiscentfruits • Legumeor pod:atwo-valved fruitformedfromasingle carpel,usuallyseveralseededandusuallyopening alongdorsalandventral sides,typicalofFabaceae. Dehiscentfruits • Lomentum:afruitderived fromasinglecarpelwhich breaksupintoone-seeded portions,asincarobtree andtamarind. Dehiscentfruits • Schizocarp:afruitderivedfroma syncarpousovarywhichbreaksupat maturityintoone-seededportions (mericarps),asinmostmembersof Lamiaceae(4mericarps)andApiaceae (2mericarps). Indehiscentfruits •Samara:adry,indehiscent,winged 1- or2-seeded(doublesamara)nut orachene,characteristicofthe genusFraxinus andAcer. •Achene: a small, dry, oneseeded, indehiscent fruit, strictly of one carpel. Indehiscentfruits • Cypsela:asmall,dry,one-seeded,indehiscentfruitformed fromtwounitedcarpels,normallyaccompainedbypappus, asintheAsteraceae.Sometimestreatedasaformofachene. Indehiscentfruits • Nut: adry,one-seeded, indehiscentfruitwithawoody pericarp(e.g.acorn,hazelnut, walnut);theseedisnotunitedto thepericarp. Indehiscentfruits • Caryopsis:adry,one-seeded,indehiscentfruit,characteristic ofgrasses(Poaceae),havingthepericarpunitedtotheseed; thegrainofacerealgrass. Fruitdispersal •Fruits of devil’s claw with hooks joining to fur coat of animals. Pharmaceuticalimportanceof fruitsandseeds •Thefollowingfruitsareusedinpharmaceutical preparations:papaya,orangeandcitrus,anise,star anise,capsicum,senna. •Severalplantsprovideseedcontainingoilstobeused inpharmaceutics:rape,linseed,cotton,castor,coconut, sesame,etc. •Otherseedusedinpharmaceuticsare:coffee,cocoa, sweetandbitteralmond,walnut,hazelnut,chestnutetc.