AnIntroductiontoBeeBiology

ByDavidStone,M.S.(Entomology) BiologyTeacher UniversityLaboratoryHighSchool Urbana,Illinois June,2005

PreparedfortheUIUCBeeSpaceProject www.beespace.uiuc.edu

Part1:OverviewofBees
Thereareanestimated30,000beespeciesworldwide.Thevastmajorityofthesespecies aresolitaryanddonotproducehoneyorlargenestswithyoung,andthereforedonot exhibitcolonydefense(Buchmann,2005). Whenonetypicallythinksofabee,thespeciesthattypicallycomestomindisthe westernhoneybee,Apismellifera.ThegenusApisiscomprisedofeightspecies.Apis melliferaiscomprisedof24differentraces.Themostcommoncommercialproduction raceisApismelliferaligustica,commonlyreferredtoasItalians.Thisraceisknownfor itshighrateofhoneyproductionanditsgentlenature,makingitafavoriteinapiaries,or commercialbeeproductionfacilities.

Italianbeeworkinginitshive.PhotobyZachHuang.

Themajorityofbeesthatoneseesoutsideofahiveareworkers(sterilefemales).A typicalhoneybeecolonyconsistsof50,00060,000sterileworkers,500to1000drones (fertilemales)andonequeen,theonlyfertilefemaleinthecolonyandmotherofthe entirepopulationofthehive(Bishop,2005). Somepeopleconfusebeeswithwasps.Beestendtobevegetariansandaregenerally hairy,whereaswaspstendtobecarnivorousandhairless(Buchmann,2005).Thevast majorityofBeeSpaceactivitiescenteraroundthewesternhoneybee,Apismellifera.

Beesorwasps?TheseEuropeanPaperWaspsshowthe hairlessbodiestypicalofwasps.PhotobyZachHuang.

Part2:BeeAnatomy
Likeallinsects,thebodyofabeeconsistsofthreeregions,thehead,thethorax,andthe abdomen. Theheadhousestwocompoundeyes,whichareusedfordistancevisionoutsideofthe hive,aswellasorientingthebee'sflightrelativetothesun.Eacheyeconsistsof3000to 5000visualprocessingunitscalledommatidia.Theeyesdonotperceiveshapesclearly butidentifycolorwell.Abee'scompoundeyesarereceptivetoultravioletlight,butless receptivetoreds.Beesrecognizeblue,yellow,whiteandblack.

Workerheadshowingdistinctcompoundeyes,antennae,andchewinglappingmouthparts.PhotobyZachHuang.

Youcanexperiencethevisualworldofthebee,seeingtheworldastheydo,atBEYE: Theworldthroughtheeyesofabee. Simpleeyes,calledocelli,arefoundnearthefrontandtopofthehead.Ocelliregister intensity,wavelength,anddurationoflight.Atdusktheocelliestimateextentof approachingdarkness,causingthebeestoreturntotheirhives.

Topofthehead,showingthethreeocelli.PhotobyZachHuang.

Antennaereceiveandanalyzehighlyvolatilesubstancesthatareresponsibleforodorand taste.Antennaealsoperceivevibrationsandmovementofair,sounds,temperature(the 2

fiveterminalsegmentsoftheflagellum)andhumidity(theeightterminalsegmentsofthe flagellum).

Workerantennashowingitsthreeregions.PhotobyZachHuang.

Thethoraxincludesthelegsandthewings.Attheendofeachlegarestructurescalled tarsi,whichtastewhattheytouch(morespecifically,theydetectqualityand concentrationofdifferentchemicals).Clawsandarolia(softpadsbetweenthepaired clawsofeachleg)combinetoprovideaneffortlessholdonbothsmoothandrough surfaces(Apiculture:KnowtheBee,ManagetheApiary).

Pairedclawsandsuctionpadlikearoliumattheendofabee'sleg.PhotobyZachHuang.

Thefirst(frontmost)pairoflegshasanotchinitsfirstterminalsegmentforcleaning antennae.Themiddlepairhasspinesononesidespecializedforremovalofmassesof pollenbroughttothehive.Thethird(hindmost)pairoflegseachpossessapollenbasket (corbicula)in whichthepollenmassiskeptduringtransportationfromtheflowerstothe hive.Thelowersideofthispairoflegsalsopossessesarowofstiffhairs,collectively calledthepollencomb.

Frontlegshowingantennacleaningnotch.

Hindlegshowingpollenbasketandcomb.

Wingsofeachbeespeciesvaryintheirvenation(vein)pattern.Theslightdifferencesin Apismelliferawingvenationcanbeusefulindifferentiatingbetweenraces.Theforewing isalwayslargerthanthehindwing.Thefrontandhindwingsareheldtogether(coupled) byapproximately20smallhookslocatedalongthefrontmarginofthehindwing.Bee wingscanbeatnearly200timespersecond.

Coupledforeandhindwings.PhotobyZachHuang.

Theabdomenconsistsofsevenvisiblesegments.Thefirstisverynarrowedandmakes upthepetiole(waist)ofthebee,whiletheseventhsegmentofworkers(sterilefemales) andqueensincludesthesting(Apiculture:KnowtheBee,ManagetheApiary).Wax glandsontheundersideofworkerabdomenssecretethewaxthatmakesupthe honeycomb.

Waxscalessecretedbythefourpairsofglandsontheundersideofaworker'sabdomen.PhotobyZachHuang.

Thestingisamodifiedovipositor,soitisfoundonlyinfemales.Whenpushedfromthe endoftheabdomen,itlocksintopositionatarightangletothebase.Muscular abdominalplatesthenpushthestingerintotheflesh.Thestinghasascalpelsharppoint, withtwoserratedretractablerods(lancets)onthesides.Thevenombulbispositionedat thetopofthesting.Itcontinuestopumpvenom30to60secondsafterbreakingofffrom theabdomenoftheworkerbee.

Stingwithattachedvenomsack.PhotobyZachHuang.

Closeupofstingshowingbarbsthatpreventeasyremoval.Photoby ZachHuang.

Uptohalfofthevenomstoredinthebulbconsistsofmelittin,achemicalsubstancethat causespain,impactsbloodvessels,anddamagestissues.Inresponse,thebodyofthe stungorganismproduceshistamines,whichcauselocalizeditching,rednessandswelling. PhotolipaseA2andhyaluronidasecontributetotheswellingandspreadofthetoxin. Additionallyalarmpheromoneisreleasedatthetimeofthesting,stimulatingfurther defensiveresponseintheworkers.Eachworkerdiesshortlyafterstinginghervictim becausethestingandpartofthedigestivetractareleftareleftatthesiteofthestinging incident(Bishop,2005).

Tearingawayofstingandpartofdigestivetract.PhotobyZachHuang.

Part3:BeeLifeStages
Typicalofthemostadvancedinsects,beesexhibitcompletedevelopmentorcomplete metamorphosis.Thismeansthattheyoungandtheadultslookverydifferentandthediet oftheyoungandtheadultstypicallydiffer,preventingtheparentsfromcompetingwith theiroffspringforresources.Thelifestagesareegg,larva,pupaandadult.Development fromeggtonewworkertypicallytakestwo(TalesFromtheHive,2000)tothreeweeks (Bishop,2005). Egg Theeggsaresometimesdescribedashavinganappearancesimilartosausageshaped poppyseeds.Eachegghasasmallopeningatthebroadendoftheegg,themicropyle, thatallowsforpassageofsperm.Hatchingtakesplacethreedaysafteregglaying(Jean Prost,1994). Larva Thelarvalstagelastseighttoninedays.Uponhatching,thelarvaisalmostmicroscopic, resemblingasmall,white,curved,segmentedwormlackinglegsandeyes.Forthefirst twodays,alllarvaearefedadietofroyaljelly.Beginningthethirdday,workerlarvae arefedhoney,pollenandwater,whilethelarvaedestinedtobecomequeenscontinueto receiveroyaljellythroughouttheirlarvallives.Regardlessofwhetherthelarvaismaleor female,itmoltsfivetimesduringitslarvalstage(JeanProst,1994).

Workerscaringforlarvae.PhotobyZachHuang.

Careofthelarvaeisconstant.Eachlarvareceivesanestimated10,000mealsduringthis stage.Larvalweightincreases51/2xduringthefirstday,1500xinsixdays(TalesFrom theHive,2000). Larvalstagedurationsvary: 5.5daysforqueens(fertilefemales), 6daysforworkers(sterilefemales),and 6.5daysfordrones(fertilemales)(JeanProst,1994). Pupa Thepupalstageisastageofmassivereorganizationoftissues.Organsundergoa completereorganization,whilebodychangesfromthewormlikelarvalbodyshapetothe adultbodyshapewiththreedistinctbodyregions.Pupationperiodsvary:queensrequire upto7.5days,dronesrequire14.5days,whileworkersrequire12days(JeanProst, 1994).

Queenandworkerpupae.Notethelargersizeofthequeenpupa.PhotobyZachHuang.

Adult Adultbeesareeitherworkers(sterilefemales),queens(fertilefemales),ordrones(fertile males).Atypicalhoneybeecolonyconsistsof50,00060,000sterileworkers,500to1000 drones(fertilemales)andonequeen,theonlyfertilefemaleinthecolonyandmotherof theentirepopulationofthehive(Bishop,2005).

Worker(left),drone(middle)andmaturequeen(right).PhotobyZachHuang.

Workersprovidevirtuallyalloftheeffortsrequiredtomaintainfunctionwithinahive. Duringthelatterpartoftheirlife,eachwilltraveluptotwomilesinsearchofpollen, nectarandwater.Eachworkertypicallygoesontenfoodgatheringjourneysperday, eachlastingapproximatelyonehour.Thisheavyworkloadtakesitstolleachworker livesforaboutamonthpriortowearingout(TalesFromtheHive,2000). Immediatelyafteremergingfromitspupalcocoonwithinoneofthemanybroodcells,it immediatelygoestowork.Duringthefirstfourdaysofitsadultlife,eachworkeris cleanedandfedbytheotherbeeswhileitsbodyhardensanditbeginstoproduce substancesinvariousglands.Activitiesduringthenextseventeendaysincludecleaning, feedinglarvae,manipulatingwax,processinghoney,guarddutyandairconditioningthe hivebyfanning.Anyoftheseactivitiescanbedoneatanytimebasedontheneedsofthe colony.

Workerscaringforaqueenlarvawithinitsqueencell.PhotobyZachHuang.

Onday21theworkerleavesthehive,andworksforanother20days,bringinginpollen, nectar,water,andpropolisbeforetakingitsfinalflightawayfromthehiveanddying (Hooper,1976). Pollen,aplantproteinsourcefortheyoung,providesnitrogen,phosphorus,aminoacids, andvitaminsessentialfordevelopmentofthesevegetarians.Polleniscollectedinpollen baskets(corbicula)ontheworkers'rearlegs. Nectar,obtainedfromfloralnectariesdeepwithinflowers,providesapurecarbohydrate sourceforallstages.Eachworkersfillsherhoneysacwithinherdigestivesystem, increasingherweightbyuptoonehalf.Uponarrivalatthehive,theworkerregurgitates thecontentsofthehoneysactotheyoungerworkerswithinthehive.Theseyounger workersreceivethenectar,whichisprocessedbyenzymeswithintheirhoneysacs,and tippedintostoragecellswhereitripensforfivedays.Atthispointthesubstancebecomes honey,andthecellcontainingitiscappedforstorage.Nectarfrom5millionflowersis requiredtoproduceasinglepintofhoney(TalesFromtheHive,2000). Waterisessentialforhydratingalloftheindividualswithinahiveandcoolingit throughouttheyear.Approximatelyfivegallonsarerequiredtohydrateandcoolthe colonyeachyear(Hooper,1976). Propolis,thefinalsubstancebroughtintothehive,is alsocalledbeeglue.Itisaplantresinusedtobuildandmaintainhives. Queenscanbedistinguishedfromworkersbytheirlongertaperedabdomensandgreater size.Queenshavethelongestlifespanofallofthebeeswithinthehive.Theirmajorrole centersaroundegglayingtoinsurethevastnumbersofindividualsrequiredtomaintaina hive.

Maturequeenwitheggs.PhotobyZachHuang.

Colonieswillmakeanewqueeniftheoriginalisailingorinfertile.Thisisdoneby producingaspecialwax cellaround7or8fertilizedeggs,theoblongarmoredincubator lookssomewhatlikeapeanut.Eggsandlarvaeareslatheredwithroyaljelly(vitaminrich hormonalgoomadebyworkers)foratwoweekperiod,afterwhichanewqueen 9

emerges.Thefirstnewqueentoemergestingsallhersisterswithinthespecializedwax cell(allofwhomarepotentialqueens)andmaykilltheoriginalqueen(hermother). Fivetofifteendaysafteremergencefromherpupalcocoonandcell,theyoungqueen fliesoff,matingwithasmanyastendronesoveraseveraldayperiod(JeanPropst, 1994).Shewillstorethespermfromthesematingsinaspermathecaforthedurationof herlife,nevertomateagain.

Spermathecastoringspermfrommatings.Spermmaybeviableuptofouryears.PhotobyZachHuang.

Shereturnstothehiveandbeginslayingupto1,500eggsperday.Queenstypicallylay severalhundredthousandeggsovertheirlifetime(TalesFromtheHive,2000).Aftertwo tofouryears,thequeenusesupallofherstoredspermandbeginsproducingunfertilized eggs,whichgiverisetodrones.Usuallytheworkersraiseoneormorequeensfromthe lastofthefertilizedeggstoreplacethenewqueen(JeanPropst,1994).Tomaximizehive productivity,honeyfarmersreplacethequeenannuallyoreveryotheryear(Bishop, 2005). Dronesarethemalebeeswithinacolony.Dronescanbedistinguishedfromworkersand queensbytheirlargesize,rectangularabdomens,largeconspicuouseyes,andnoisy flight.Alldroneslackasting,andhavemoreeyefacetsthanaworker(6,0007,000vs. 3,0005,000).

Worker(left)anddronewithreproductivestructuresprotruding.PhotobyZachHuang.

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Dronesresultfromunfertilizedeggs.Theyemerge24daysaftertheeggislaid.Drones arecapableofextractinghoneyfourdaysafteremergence,butprefertobefedby workers.Unlikeworkers(sterilefemales),dronescan'tflywell,don'tgatherfoodforthe colony,don'tclean,don'tsecretewax,anddonotcareforyoung.Theroleofthedronesis largelytofertilizenewqueens.Agroupofdronesfollowseachvirginqueenonherearly flights.Severalmaleswillmatewitheachvirginqueenwhileflying,dyingimmediately aftermatingsincehisreproductiveorgansandtheendofhisabdomenbreakoff, temporarilypluggingtheendofthequeen'sreproductivetractandabdomen Assumingallgoeswell,dronestypicallyliveforabout50days.Ifthereisafertilefemale inresidence,theworkersmaywithholdfoodfromthedronesorgnawoffthedrones' wingsandlegs.Byfall,allofthemalesandmalelarvaeareevictedfromeachcolony (Bishop,2005).

Part4:AfricanizedBees(KillerBees)
Inthe1950s,BrazilianstriedtojumpstarttheirhoneyindustryusingEuropeanbees. HowevertheEuropeanbeesdidn'tadaptwelltotheBrazilianclimate.Africanbees, muchbetteradaptedtotheenvironmentalextremesofmanyoftheareasinterestedin commercialbeekeeping,wereimportedinhopethattheywouldbecomeacommercially viablespecies.Africanbeestendtobe10%smallerthanEuropeanbees.Additionally, theyare25%lighter,theyreproduceearlier,andtheyhaveashorterlifespan.

Africanizedbeesattheentranceofahive.PhotobyZachHuang.

Atthetime,theprospectivebeekeeperswereunawareofthefactthatAfricanbeestendto bemuchmorefierce,dominant,andaggressivethantheEuropeanbees.Inaddition,

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Africanbeestendtobenomadic,whichmeantthattheyreadilyleftmanmadehives, movedintothejunglesandestablishingtheirowncoloniesawayfromhumanhabitation. Behaviorally,Africanbeestendtobeoffensive,ratherthandefensive,whenother organismscomeintocloseproximityoftheirhives.Infact,thedefensiveresponseofa singlebeecausesarapidresponsesothatsuddenlyandseverelytheorganismmay experienceattackbyhundredsofbees.Africanizedbeesmaypursueinvadersforhours aftertheorganismunderattackhaslefttheimmediatehivearea.Thoughthevenomis virtuallyidenticaltothatofEuropeanbees,theintensityoftheattackandthevolumeof venomthattheinvadingorganismexperiencesledtothepopularpressdesignating Africanizedbeesas"killerbees." TheseaggressivebeesrapidlytookoverEuropeanbeecolonieswiththreeyearsoftheir initialBrazilianintroduction.IncontrasttotheEuropeanbees,Africanizedbeehoney harvestisunpleasantandmaybedangerous,whiletheyieldisunreliable. AfricanizedbeesexpandedtheirrangetoVenezuelain1973,Mexicoin1986,and establishedsomecoloniesintheU.S.by1990.TheynowcanbefoundinNewMexico, Arizona,Nevada,andCalifornia(Bishop,2005).

ForagingAfricanizedBee(Tucson,Arizona).PhotobyZachHuang.

Part5:BeeParasites/Competitors
Beeshaveseveralparasites/competitors.Thespeciesmakingthegreatestimpacton honeybeepopulationsincludebacteriaandmites.Eachorganismanditsimpactsare brieflyoutlinedbelow:

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Bacteria Parasiticbacteriamaketheirmostsignificantimpactonyounglarvae.Thediseasesmost commonlycausedbythesebacteriaareknownasAmericanfoulbroodandEuropean foulbrood. Americanfoulbroodiscausedby Bacilluslarvae.Infectedcoloniescontainingbrood (larvaeandpupae)oftendie.Becausethebacterialsporesremainviablefordecades,itis essentialthatmeasuresbetakentopreventinfectioninthefirstplace.Sporesare transmittedtothelarvaebyadultworkers,whocleanandfeedthelarvae.Nursebees oftenremovesomeinfectedlarvaethoughthedecomposedlarvalremains,whichare typicallyloadedwithspores,preventtheworkersfrombeingabletoeffectivelycleanout thebroodcellscontainingtheinfectedanddeadlarvae.

LarvakilledbyAmericanfoulbrood.Thisdeadlarvacontainsthousandsofsporesthatwillbespread throughoutthecolonybytheworkersengagedinnursingactivities.PhotobyZachHuang.

Ifthelarvalcellofaninfectedlarvaiscappedpriortodeath,thecappingoverthelarva becomesmoist,darkcolored,andsunken.

Broodcellsexhibitingthebrown,sunkencapscharacteristicofcolonyinfectedwithAmericanfoulbrood. PhotobyZachHuang.

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Thisdiseaseishighlycontagious.Destructionof infectedcoloniesbyburningistheonly properwaytocontrolthespreadofthisdisease.Theincidenceofthisdiseaseisestimated tobe20%intheU.S. Europeanfoulbroodiscausedby Melissococcuspluton.Itaffectslarvaeupto48hours old,andkillsthemwithinseveraldays.UnlikeAmericanfoulbrood,itsoccurrenceis cyclic,witharapidincreaseearlyinthespring.Thegreatestimpactisonsmallcolonies. ThisconditionislessofaconcernthanAmericanfoulbrood infactmanyhiveowners neverrealizethattheircoloniesareinfected.Coloniesmovedaboutforpollinationand thosewithlowfoodreservesaremostlikelytobeimpacted.Successfultreatmentofthis conditionoftenoccursasaresultoffeedingthebeesantibiotics(Crane,1990).

LarvaewithEuropeanfoulbrood.Notetheyellowstreakswhichbroadenandbecomebrownaslarvaltissueis destroyed.Thiscontinuesuntilthelarvadies.PhotobyZachHuang.

Mites VarroaMites Varroamitesarenaturalparasitesoftheeasternhoneybee,Apissurana,fromsoutheast Asia.UnfortunatelythehoneybeesthatarefoundintheU.S.(Apismellifera)haveno adaptationsthatprovidethemany significantdegreeofprotectionfromtheseparasitic mites(Buchmann,2005).Varroamitesappearasbrownspecksrangingfrom12mm.in diameter.Malesareroundandapproximately1mm.indiameter,whereastheoval femalesapproach2mm.indiameter.

Variousstagesofvarroamitedevelopment.

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Earlyinthe1980svarroamitesbegantomakeaseriousimpactoncommercialhives. Thesemitesfeedonlarvaeandpupae,feedingpreferentiallyondronelarvaeandpupae.

Larvacoveredwithfivevarroamites.PhotobyZachHuang.

Inseriouscases,thepupaearekilledordevelopintoadultsexhibitingreducedsize, incompletewingdevelopment,andatrophiedlegs.Oneof themajorproblemsindealing withtheseorganismsisthattheiractivitiesgolargelyunnoticedwithincappedcells, awayfromtheviewofeventhemostvigilantbeekeepers.Adultsbeesbecomeexhausted intheireffortstoremovemites.Feedingofthemitescausestheadulttolosevitalbody fluids,whilethebreaksintheexoskeletoncausedbymitefeedingleadtoincreased likelihoodofinfectionbybacteria,viruses,andotherpathogenicagents.Oftenthemites donotappeartomakeasignificantimpactonthebeecolonyuntilthethirdyearofmite populationestablishment,whenthemitesmayconsumetheentireyear'slarvaeand pupae.Acaracides,pesticidesdevelopedspecificallyformitecontrol,mayreducemite numbersthoughthereisnocompletely effectivecontrolmethod(JeanPropst,1995). Halfofthepopulationsofsomebeeshavedecreasedoverthepast10yearsasaresultof thismite(Buchmann,2005).

Varroamitesonthethoraxofaworker.Notetheincompletewingdevelopmentinthebottommostadult.Thisis likelytheresultofvarroamitetissuedamageduringthepupalstage.PhotobyZachHuang.

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TrachealMites Thesemitesareanaturalparasiteofthewesternhoneybee.TheyhavebeenintheU.S. since1984,andaregenerallyconsideredarelativelyminorpestinmanypartsoftheU.S. (Buchmann,2005).Onlyyoungadultbeesaresusceptibletothismite.Thesemitesenter thefirstorsecondspiracles(airholes)andfeedonthewallofthetrachea,thetubesused forgastransport.Trachealmiteimpactoncoloniesisoftenminimal,thoughinfestations incolonieswithinsufficientfoodresourcesfurtherweakensthecolony,andmay contributetotheeventualdeathofthethesecolonies(Crane,1990).

Twotrachealmiteswithinthetracheaofaparasitizedworker.PhotobyZachHuang.

Part6:BeeProducts
Themajorityofcommercialapiaryprofitscomefrompaymentforpollinationservices.In additiontopollinationprofits,severalbeeproductsareeconomicallysignificant.Thetwo mostcommonbeeproducts,honeyandwax,areproducedandusedthroughouttheworld. Honey Honeyisproducedfromnectar,whichisobtainedfromfloralnectariesdeepwithin flowers.Nectarprovidesapurecarbohydratesourceforallstageofthebeelifecycle. Eachworkerfillsherhoneysacwithinherdigestivesystem,increasingherweightbyup toonehalf.Uponarrivalatthehive,theworkerregurgitatesthecontentsofthehoneysac totheyoungerworkerswithinthehive.

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Foragingbeeregurgitatingstomachcontentstoyoungernursebees.PhotobyZachHuang.

Theseyoungerworkersreceivethenectar,whichisprocessedbyenzymeswithintheir honeysacs,andtippedintostoragecellswhereitthenripensforfivedays.Atthispoint thesubstancehasbecomeshoney,andthecellcontainingitiscappedforstorage.

Honeyinthecells Whenthewatercontentislessthan18%,workerssealeachcellwithacap.Photo byZachHuang.

Nectarfromfivemillionflowersisrequiredtoproduceasinglepintofhoney(Tales FromtheHive,2000).Duringitslifetime,thetypicalworkermakes1/12ofateaspoonof honey.Asinglepoundofhoneyrequirestwomillionflowervisits.Theaverage commercialbeehiveproducesinexcessof160poundsofhoneyperyear(Bishop,2005). Honeyhasbeenusedasasweetenersinceancienttimes.Thecolorandflavorofhoneyis impactedbytheflowersource.Ingeneral,thedarkerthehoney,thestrongeritsflavor. Forexample,orangetreebasedhoneyhasacitrusodorandtaste(ambercolor,citrus tang,andorangearoma),whileeucalyptushoneyisdarklyaromaticandsomewhat medicinaltasting.Honeymadefromleatherwood,acomponentofaftershaves,hasan exotic,spicy,aftershavescent. 17

Buckwheathoney,likeotherdarkhoneys,isknownforitscharacteristicstrongflavorandhighlevelsof antioxidants.PhotobyZachHuang.

Inadditiontoitsvalueasasweetener,honeyhasanumberofmedicinalandantibacterial properties,andhasbeenusedinpainreliefandhealingofburnvictims.Thesugarsin honeynourishhealthycellsandhelpsupportthedevelopmentofnewwhitebloodcells. Honey'santioxidants,aminoacidsandvitaminsplayaroleinreducinginflammation. Theantibacterialactivityofhoneyrapidlykillsthepathogensthatcausetyphoidfever, bacterialpneumonia,strepthroatandbacterialdysentery.Infact,in1998honeywas proventobemoreeffectivethansilversulphadiazine,theantibacterialointmentmost widelyusedonburnsinhospitalsituations,inthetreatmentofburns. Wax Beewaxhastraditionallybeenusedinanumberofitemsincludingfurnitureandshoe polish,crayons,candles,lipbalms,lipstick,handcreams.Intermsofworldwide corporatesales,thefivelargestusersofbeewaxare1)cosmeticcompanies,2)candle makers,3)pharmaceuticalcompanies,4)dentistoffices,and5)chewinggumcompanies. Interestingly,beesconvertexcessfoodresourceswithintheirownbodiestowaxinstead offat.Waxglandsontheundersideofworkerabdomenssecretethewaxthatmakesup thehoneycomb.

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Waxscalessecretedbythefourpairsofglandsontheundersideofaworker'sabdomen.PhotobyZachHuang.

Productionofasinglepoundofwaxrequireseightpoundsofnectar.Thissamesingle poundofwaxwillbeusedtomake35,000broodcells.Thoughtherearelikelyfew peopleinvolvedinthecommercialproductionofbeesforthesolepurposeofwax production,theannualwaxharvestfromcommercialhivesoftenprovidesenoughmoney forbeekeeperstorefurbisholdhivesaswellasproducesomenewonesforthenextyear (Bishop,2005).

Newlyproducedwaxisacreamywhitecolor.Olderwaxtendstobecaramelcolored,whiletheoldestwaxis adarkchocolatebrown.PhotobyZachHuang.

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Part7:RecommendedBeeRelatedResources
Books
Bishop,H.(2005). Robbingthebees:abiographyofhoney,thesweetliquidgoldthat seducedtheworld.NewYork,NY:FreePress. Crane,Eva.(1990). Beesandbeekeeping:science,practiceandworldresources. Ithaca, NY:ComstockPublishingAssociates. Hooper,T.(1976). Guidetobeesandhoney.Emauss,PA:RodalePress. JeanPropst,P.(1994). Apiculture:knowthebeeandtheapiary.Andover,Hampshire, U.K.:OxfodandIBHPublishingCo. Thompkins,E.&Griffith,R.(1977). Practicalbeekeeping.Charlotte,VT:GardenWay Publishing. Winston,M.(1992). Killerbees:theAfricanizedhoneybeeintheAmericas.Cambridge, MA:HarvardUniversityPress.

OnlineAudioInterviews
NPRDianeRehmShowinterviewwithHolleyBishop,authorof Robbingthebees:a biographyofhoney,thesweetliquidgoldthatseducedtheworld.March30,2005.< http://www.wamu.org/programs/dr/05/03/30.php> ScienceFridayinterviewwithStephenBuchmann,authorof Lettersfromthehive:an intimatehistoryofbees,honey,andhumankind.June17,2005.< http://www.sciencefriday.com/pages/2005/Jun/hour2_061705.html>

Videotapes
TalesFromtheHive.Dir.HerbertHabersack.NOVA.2000.Videocassette.

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WebSites
General Beetography http://www.beetography.com/Top qualitybeerelateddigitalimagesof bees.ThankstoZachHuangforgrantinguspermissiontousemanyoftheseimageson thiswebsite. ApiservicesVirtualBeekeepingGallery, http://www.apiculture.com/_menus_us/index.htm BeeInformationontheWorldWideWeb,http://www.honeybee.com/index.htm BeeHoothebeekeepingdirectory Beekeepingbeekeepingsuppliers,apiculture, http://www.beehoo.com Beekeeper'sWebLinks, http://ourworld.compuserve.com/homepages/Beekeeping/weblinks.htm Beemaster'sBeekeepingCourseHomePage, http://www.beemaster.com/honeybee/beehome.htm BEESONLINE,http://www.beesonline.com/ TheBiologyoftheHoneybee,Apismellifera, http://koning.ecsu.ctstateu.edu/plants_human/bees/bees.html CyberBeeNet(Biology,Research,Beekeeping),http://www.cyberbee.net/ InGenBees,http://www.ingenbees.com/ NCBIHoneyBeeGenomeResources, http://www.ncbi.nlm.nih.gov/projects/genome/guide/bee/ ThePollinationHomePage,http://pollinator.com/

Regionbased,thoughlotsofgeneralinformation! AlaskaHoneyBeeHomePage,http://balder.prohosting.com/~starrier/ EasternApiculturalSocietyofNorthAmerica,Inc.,http://www.easternapiculture.org/ HoozierBuzz.com:TheOfficialWebsiteoftheIndianaStateBeekeepersAssociation, http://www.hoosierbuzz.com/

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IllinoisStateBeekeepersAssociation,http://www.isba.us/ IndianaBeekeeping,http://indianabeekeeper.goshen.edu/ InternationalBeeResearchAssociation,http://www.ibra.org.uk/shop/erol.html MAARECMidAtlanticApicultureResearchandExtensionConsortium, http://maarec.cas.psu.edu/ MidniteBee:BeekeepersHome,http://www.mainebee.com/

Images MicroAngela'sElectronMicroscopicImages, http://www.pbrc.hawaii.edu/bemf/microangela/ ScanningElectronMicrographsofInsects,http://www.bath.ac.uk/ceos/Insects4.html AScanningElectronMicroscopeAtlasoftheHoneyBee, http://gears.tucson.ars.ag.gov/beebook/bee.html

Miscellaneous AfricanizedHoneyBeesontheMove:LessonPlans, http://ag.arizona.edu/pubs/insects/ahb/ahbhome.html BEYE:Theworldthroughtheeyesofabee, http://cvs.anu.edu.au/andy/beye/beyehome.html

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