Palynology Introduction alynology ls Lhe sclence LhaL sLudles conLemporary and fossll palynomorphs lncludlng pollen spores dlnoflagellaLe cysLs acrlLarchs chlLlnozoans and scolecodonLs LogeLher wlLh parLlculaLe organlc maLLer (CM) and kerogen found ln sedlmenLary rocks and sedlmenLs 1he Lerm palynology was lnLroduced by Pyde and Wllllams ln 1944 followlng correspondence wlLh Lhe Swedlsh geologlsL AnLevs ln Lhe pages of Lhe ollen Analysls Clrcular (one of Lhe flrsL [ournals devoLed Lo pollen analysls and produced by aul Sears ln norLh Amerlca) Pyde and Wllllams chose palynology on Lhe basls of Lhe Creek words paluno meanlng Lo sprlnkle and pale meanlng dusL (and Lhus slmllar Lo Lhe LaLln word pollen) 1hey used Lhe Lerm palynology for Lhe sLudy of exLernal (morphologlcal) feaLures of maLure pollen gralns alynology ls an lnLerdlsclpllnary sclence and ls a branch of earLh sclence (geology or geologlcal sclence) and blologlcal sclence (blology) parLlcularly planL sclence (boLany) SLraLlgraphlcal palynology ls a branch of mlcropalaeonLology and paleoboLany whlch sLudles fossll palynomorphs from Lhe recambrlan Lo Lhe Polocene History Mohl (1833) classlfled Lhe pollen gralns accordlng Lo Lhelr aperLures aperLure membranes eLc Carl !ullus lrlLzsche whose paper 'ueber den ollen' publlshed ln 1837 conLrlbuLed Lo Lhe sLudy of pollen gralns wlLh lllusLraLlons and accuraLe descrlpLlons Pugo llscher (1890) descrlbed Lhe deLalls of exlne ln a number of planLs 1he modern era of pollen analysls and Lhe subsequenL developmenL of palynology sLarLs wlLh LennarL von osL's lecLure on CuaLernary deposlLs ln 1916 8oger Wodehouse's classlc 'ollen Cralns' was publlshed ln 1933 LrdLman's 'ollen Morphology and lanL 1axonomy' publlshed ln 1932 ls a monumenLal work ln palynology 1exL 8ook of ollen Analysls" of laegrl and lvesen (1964) ls anoLher lmporLanL work on prlnclples of pollen analysls 1he advenL of elecLron mlcroscopy puL new lmpeLus Lo Lhe sLudy of Lhe flner deLalls of Lhe enLlre pollen gralns and spores Brancbes alynology can be dlvlded lnLo 8aslc (lundamenLal) and Applled alynology 8aslc alynology deals wlLh pollen and spore morphology Lhe physlcal and chemlcal properLles of sporoderm as well as Lhe correlaLlon beLween palynology and cyLology lL also lncludes LheoreLlcal aspecLs of applled palynology Applled alynology deals wlLh applled aspecLs of Lhe palynology lL lncludes O alynoLaxonomy Lhe sLudy of pollen morphology wlLh relaLlon Lo Lhe planL Laxonomy O Ceo or alaeopalynology Lhe sLudy of fossll pollen gralns or spores O MellLLopalynology Lhe sLudy of pollen gralns found ln honey O Aeropalynology Lhe sLudy of frequency and dlsLrlbuLlon of spores ln Lhe alr O laLropalynology Lhe sLudy of medlcal aspecLs of palynology speclally Lhe allergles O harmacopalynology Lhe sLudy of roles of pollen gralns and spores ln drugs O Copropalynology Lhe sLudy of pollen gralns and spores ln excremenLs O lorenslc palynology appllcaLlon of pollen gralns as an ald ln crlmlnology and crlme deLecLlon
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\\ications lmporLance of pollen ln Lhe lmprovemenL of planLs was recognlzed as early as 717 8C by Assyrlan arLlsans AnclenL lndlan classlcs conLaln some reference abouL Lhe refreshlng pollen baLh of lndlan laburnum (Cassla flsLula) 1he cosmeLlc chesLs of LgypLlan queens are belleged Lo have conLalned pollen powder for faclal care ollen culL was developed by Amerlcan lndlans who consldered pollens of caLLall (1ypha) and corn as an emblem of peace happlness and prosperlLy ollens are also used as human food by some people Amerlcan lndlans use pollens of 1ypha for preparlng pollen soup or gruel lew Lrlbes of new Zealand lndla and aklsLan use 1ypha pollens as food by mlxlng wlLh waLer Lo form cakes and baked 1he medlclnal lmporLance of pollen and spores was long recognlzed MenLlon may be made of Lycopodlum spores used by Lhe anclenL Arablan physlclans Lo cure sLomach dlsorders spores of AdlanLum hlllpplnes used ln leprosy and skln dlseases pollens of 1ypha laxmanll used as asLrlngenL Cycas clrclnalls as narcoLlc 8ee collecLed pollens mlxed wlLh coffee mllk honey poLaLoes and chocolaLes are prescrlbed Lo cure many dlsorders ln chlldren Slgnlflcance of pollen ln agrlculLure has been recognlzed ln recenL Llmes SelecLlon of pollen wlLh Lhe deslred poLenLlal from deslred lndlvlduals of any one crop populaLlon ls consldered as an essenLlal prerequlslLe for planL breedlng SLorage of pollen ls as lmporLanL as sLorage of seeds ln malnLalnlng planL geneLlc resources ln many research lnsLlLuLes Lhe haplold planLs are used for Lhe malnLenance of geneLlc sLock lL ls clalmed by Chlnese and !apanese sclenLs LhaL haplold planLs of rlce (Cryza saLlva) developed from pollens have been Lransferred from LesL Lubes Lo fleld condlLlons ln some counLrles sLored pollens are dlssemlnaLed over crop flelds by means of sprayers drawn by LracLors or by breaklng ollen bombs or by hand on days wlLh moderaLely hlgh wlnds Successful polllnaLlon ensures abundanL frulL sulL and lncreased seed producLlon ollens are ldeal maLerlal for experlmenLally conLrolllng Lhe geneLlc behavlor of planLs L produce more and more useful hybrlds ollen lrradlaLlon ls used Lo meeL Lhe problems of lnLraspeclflc or lnLerspeclflc lncompaLlblllLy by lnduclng deslred muLaLlonal changes ln Lhe pollen genome lnducLlon of male sLerlllLy becomes a boon ln crosslng self polllnaLed planLs llke grapes Aeropalynologlcal sLudles of aeropollens have new vlsLa ln conLrolllng Lhe weed and has been pracLlced successfully ln Lngland SLudy of honey pollens provldes lnformaLlon abouL Lhe planLs from whlch Lhe necLar and pollens have been gaLhered Such sLudles can deLecL adulLeraLlon ln honey Sharma and nalr (1963) sLaLed LhaL lf honey conLalns 30 percenL of more pollens of one planL lL should be called unlfloral honey oLherwlse lL should be referred Lo as MulLlfloral honey alynology ls used for a dlverse range of appllcaLlons relaLed Lo many sclenLlflc dlsclpllnes O 8losLraLlgraphy and geochronology CeologlsLs use palynologlcal sLudles ln blosLraLlgraphy Lo correlaLe sLraLa and deLermlne Lhe relaLlve age of a glven bed horlzon formaLlon or sLraLlgraphlcal sequence O alaeoecology and cllmaLe change alynology can be used Lo reconsLrucL pasL vegeLaLlon (land planLs) and marlne and freshwaLer phyLoplankLon communlLles and so lnfer pasL envlronmenLal (palaeoenvlronmenLal) and palaeocllmaLlc condlLlons O Crganlc palynofacles sLudles whlch examlne Lhe preservaLlon of Lhe parLlculaLe organlc maLLer and palynomorphs provldes lnformaLlon on Lhe deposlLlonal envlronmenL of sedlmenLs and deposlLlonal palaeoenvlronmenLs of sedlmenLary rocks
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O CeoLhermal alLeraLlon sLudles examlne Lhe colour of palynomorphs exLracLed from rocks Lo glve Lhe Lhermal alLeraLlon and maLuraLlon of sedlmenLary sequences whlch provldes esLlmaLes of maxlmum palaeoLemperaLures O Llmnology sLudles lreshwaLer palynomorphs and anlmal and planL fragmenLs lncludlng Lhe praslnophyLes and desmlds (green algae) can be used Lo sLudy pasL lake levels and long Lerm cllmaLe change O 1axonomy and evoluLlonary sLudles O lorenslc palynology Lhe sLudy of pollen and oLher palynomorphs for evldence aL a crlme scene O Allergy sLudles SLudles of Lhe geographlc dlsLrlbuLlon and seasonal producLlon of pollen can help sufferers of allergles such as hay fever O Mellssopalynology Lhe sLudy of pollen and spores found ln honey O Archaeologlcal alynology examlnes human uses of planLs ln Lhe pasL 1hls can help deLermlne seasonallLy of slLe occupaLlon presence or absence of agrlculLural pracLlces or producLs and planLrelaLed acLlvlLy areas wlLhln an archaeologlcal conLexL 8onflre ShelLer ls one such example of Lhls appllcaLlon 8ecause Lhe dlsLrlbuLlon of acrlLarchs chlLlnozoans dlnoflagellaLe cysLs pollen and spores provldes evldence of sLraLlgraphlcal correlaLlon Lhrough blosLraLlgraphy and palaeoenvlronmenLal reconsLrucLlon one common and lucraLlve appllcaLlon of palynology ls ln oll and gas exploraLlon alynology also allows sclenLlsLs Lo lnfer Lhe cllmaLlc condlLlons from Lhe vegeLaLlon presenL ln an area Lhousands or mllllons of years ago 1hls ls a fundamenLal parL of research lnLo cllmaLe change etbods of Study lor Lhe sLudy of Lhelr exLernal morphology pollen slldes are prepared by aceLolysls as suggesLed by LrdLman (1932) lolyoomotpbs ote btooJly JefloeJ os otqoolcwolleJ mlctofosslls betweeo 5 ooJ 500 mlctomettes lo slze 1bey ote exttocteJ ftom tocks ooJ seJlmeots botb pbyslcolly by wet slevloq ofteo oftet olttosoolc tteotmeot ooJ cbemlcolly by osloq cbemlcol Jlqestloo to temove tbe ooootqoolc ftoctloo lot exomple polyoomotpbs moy be exttocteJ osloq byJtocblotlc oclJ (ncl) to Jlqest cotbooote mloetols ooJ byJtoflootlc oclJ (nl) to Jlqest slllcote mloetols lo soltoble fome copbootJs lo speclollst lobotototles Samples are Lhen mounLed on mlcroscope slldes and examlned uslng llghL mlcroscopy or scannlng elecLron mlcroscopy Cnce Lhe pollen gralns have been ldenLlfled Lhey can be ploLLed on a pollen dlagram whlch ls Lhen used for lnLerpreLaLlon ollen dlagrams are useful ln glvlng evldence of pasL human acLlvlLy (anLhropogenlc lmpacL) vegeLaLlon hlsLory and cllmaLlc hlsLory alynology uses many Lechnlques from oLher relaLed flelds such as geology boLany paleonLology archaeology pedology and geography oen or\boogy ollen morphology deals malnly wlLh Lhe walls of pollen gralns and spores and noL wlLh Lhelr llvlng conLenLs 1he pollen wall has a very complex sLrucLure LhaL ls reflecLlve of LhaL speclflc specles funcLlonal adapLaLlons 1he exlne ls Lhe ouLer layer of a llvlng pollen graln lL ls composed of sporopollenln wlLh small quanLlLles of
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polysaccharldes Sporopollenln ls very chemlcally sLable and lL ls reslsLanL Lo almosL all klnds of envlronmenLal damage lL ls equlpped wlLh aperLures lL lL dlvlded lnLo Lwo sub layers Lhe ouLermosL sexlne and Lhe unsculpLured underlylng nexlne 1he sexlne has surfaces LhaL are sculpLured ln elaboraLe ways wlLh reLlculaLely arranged perforaLlons 1hese glve Lhe exlne and amorphous or granular appearance 1he lnner layer of a llvlng pollen graln ls called Lhe lnLlne lL ls composed of cellulose and ls very slmllar ln consLrucLlon Lo ordlnary planL cell walls A layer called Lhe endexlne separaLes Lhe sexlne and lnLlne 1he endexlne has a lamlnaLed appearance ollen gralns are generally classlfled accordlng Lo Lhelr physlcal appearance 1here are Lhree crlLerla of classlflcaLlon 1) Lhe number and poslLlon of Lhe aperLures 2) Lhe shape of Lhe pollen graln as a whole and 3) Lhe flne elaboraLe sLrucLure on Lhe sexlne AperLures are any mlsslng parLs of Lhe exlne whlch are lndependenL of Lhe exlne paLLern AperLures are blg and Lhey cuL across Lhe flne sLrucLure paLLern on Lhe surface of Lhe pollen graln 1here are Lwo Lypes of aperLures porl or pores are mosLly lsodlameLrlc aperLures alLhough Lhe can be sllghLly elongaLed wlLh rounded ends colpl or furrows are long and boaL shaped wlLh polnLed ends Colpl are LhoughL Lo be more prlmlLlve ln llvlng pollen gralns Lhese aperLures are noL acLually open lnsLead a very Lhln layer of exlne covers Lhem Cralns wlLh porl are called poraLe Lhose wlLh culpl are called culpaLe and Lhose wlLh boLh porl and culpl are called colporaLe lf Lhelr aperLures are arranged equldlsLanLly around Lhe equaLor of Lhe pollen gralns Lhey are asslgned Lhe preflx zono lf Lhey are scaLLered all over Lhe surface of Lhe pollen graln Lhey are asslgned Lhe preflx panLo 1he number of aperLures ls also lndlcaLed by preflxes mono for one aperLure dl for Lwo aperLures Lrl for Lhree aperLures and so on
1he shape of a pollen graln refers Lo Lhe shape of Lhelr ouLllne ln polar and equaLorlal vlews 1he shape of a graln can someLlmes be useful ln ldenLlfylng of pollen specles buL noL usually 1he sculpLure refers Lo Lhe flne sLrucLure and paLLern of Lhe sexlne lL ls composed of small radlally dlrecLed rods lf Lhese rods supporL someLhlng (such as a plaLe or a small knob) Lhey are called columnellae lf Lhey do noL supporL anyLhlng Lhey are called bacula 1he shape of Lhe rods can furLher classlfy Lhem lf Lhey are club shaped Lhey are called clavae lf Lhey are sharply polnLed Lhey are called echlnae lf Lhey have swollen heads Lhey are called plla and lf Lhey are shorL and globular Lhey are called gemmae 1here are many more classlflcaLlons for Lhe shape of rods on Lhe surface of Lhe sexlne buL Lhese four are Lhe mosL common ln general characLers of pollens are sLudled under Lhe dlvlslons pollen unlLs polarlLy symmeLry aperLures slze and shape exlne sLraLlflcaLlon and exlne palynogram oen Units Slngle pollen gralns and spores whlch when maLure do noL remaln unlLed wlLh oLher polleng ralns and spores whlch are referred Lo as monads lf pollen gralns are unlLed ln palrs Lhey are called dyas (Scheuchzerla palusLrls podosLemonaceae) A LeLrad conslsLs of four pollen gralns An ocLad of elghL a polyad of more Lhan elghL pollen gralns are also found 1eLrads can be caLegorlzed as follows O 1eLrahedral one of polleng ralsn lylng cenLrally over Lhe oLher Lhree (urlmys) O 1eLragonal (lsobllaLeral) all pollen gralns ln one plane blunLly quadrangular (MonocoLyledons) O 8homboldal (Anona murlcaLa) O uecussaLe (Cross) pollen gralns palrwlse aL rlghL angles Lo each oLher (Magnolla randlflora)
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O 1shaped Lhe flrsL dlvlslon of Lhe MC ls ransverse and a dyad cell ls formed elLher Lhe upper or Lhe lower dyad cell undergles verLlcal or longlLudlnal Lo yleld lnverLed 1shaped conflguraLlon (ArlsLolochla) O Llnear Lhe flrsL dlvlslon ls always Lransverse and glves rlse Lo dyad cells and second dlvlslon also Lransverse Lo glve llnear arrangemenL (Mlmosa pudlca)
ln sedges (cyperaceae) monad llke LeLrads are formed wlLhouL dlvldlng walls beLween Lhe four componenLs Cne nucleus ls normally developed whereas Lhe remalnlng Lhree are obllLeraLed Such Lype of degraded LeLrad ls called pseudomonad ln Mlmosaceae each of Lhe Lrad cells dlvldes once or Lwlce and Lhese arlses a massulae of elghL Lo 64 cells whlch remaln LogeLher afLer maLurlLy (buL noL Lhe enLlre producL of one Lheca) 1hey consLlLuLe a small unlLs and also called polyads
ln polllnlum a unlL of all gralns produced by one Lheca (locule lnslde Lhe anLher) aggregaLe LogeLher Lo form one slngle unlL An enLlre pollen mass of a Lhecal chamber form Lhe polllnlum 1he polllnlum apparaLus ls Lhe funcLlonal unlL of a corpusculum wlLh lLs Lwo aLLached arms and polllnla (CaloLropls Crchldaceae) oarity All pollen gralns have a LeLrad sLage durlng developmenL lL ls aL Lhls sLage LhaL polarlLy ls deLermlned AL Lhe LeLrad sLage Lhe end of a pollen dlrecLed Lowards Lhe cenLre of LeLrad ls called proxlmal pole whlle opposlLe ls Lhe dlsLal pole 1he dlsal pole faces perlpheral slde 1he polar axls ls always perpendlcular wlLh Lhe dlsLal pole aL Lhe ape and Lhe proxlmal pole aL Lhe base 1he cenLre of each paL ls called pole and Lhe lmaglnary sLralghL llne Lhrough Lhe poles ls called Lhe polar axls and each plane Lhrough Lhls axls ls a polar plane 1he border llne beLween Lhe dlsLanL parL and Lhe proxlmal parL of Lhe polen graln ls Lhe equaLor and Lhe plane Lhrough Lhe equaLor ls called Lhe equaLorlal plane
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ollen gralns may be O Apolar pollen gralns wlLhouL dlsLlncL polarlLy for example spheroldal gralns LhaL ls wlLhouL dlscernlble poles once Lhe gralns have been separaLed from Lhe pollen LeLrads O olar pollen gralns wlLh recognlzable poles even afLer Lhe gralns have been separaLed from Lhe pollen LeLrad ln lsopolar gralns Lhe dlsLal and proxlmal poles look 'allke' ln heLeropolar Lhe Lwo faces are essenLlally dlfferenL Cralns whlch have more or less curved equaLorlal plane and possess only sllghL dlfferences beLween Lhe dlsLal and proxlmal faces (one conves and oLher concave) may be Lermed paralsopolar (sublsopolar) ln llnear LeLrad Lhere ls no polarlLy ln Lhe usual sense of Lhe Lerm Symmetry lL ls Lhe quallLy lnherenL ln a body whlch ls capable of dlvlslon lnLo slmllar or equal halves Spores may be symmeLry belng bllaLeral or radlal ln radlosymmeLrlc spores Lhe shape ls such LhaL any plane lncludlng Lhe polar axls LhaL posses Lhrough wlll produce ldenLlcal halves 8llaLeral spores have Lwo verLlcal planes of symmeLry Some spores are asymmeLrlc (wlLh no symmeLry aL all) Some spores possess flxed shape (flxlform) whlle some are wlLhouL a flxed shape (nonflxlform) Sba\e Accordlng Lo LrdLman radlally symmeLrlcal pollen gralns are shaped llke a roLaLlon elllpsold wlLh Lhe polar axls as Lhe axls of roLaLlon 1he shape ls calculaLed by Laklng measuremenLs of polar axls () and Lhe maxlmum breadLh (L) ln Lhe equaLorlal vlew of Lhe graln and applylng Lhe formula /L x 100 1he vale obLalned glves Lhe shape 1he shape classes are as follows Shape C|asses ]L k 100 erob|ate 30 Cb|ate 30 73 Subsphero|da| 73 133 Subob|ate 73 88 Cb|ate sphero|da| 88 100 ro|ate sphero|da| 100 114 Subpro|ate 114 133 ro|ate 133 200 erpro|ate 200
Confervold gralns (flllform fllamenLous) are descrlbed as long slmple one walled Lhreads arranged ln parallel masses equallylng Lhe anLher ln lengLh and Lendlng Lo becoe very Langled when dlsLurbed (ZosLera sp)