Palynology

Introduction
Palynology is the science that studies contemporary and fossil palynomorphs, including pollen, spores, dinoflagellatecysts,acritarchs,chitinozoansandscolecodonts,togetherwithparticulateorganicmatter(POM)and kerogenfoundinsedimentaryrocksandsediments. ThetermpalynologywasintroducedbyHydeandWilliamsin1944,followingcorrespondencewiththeSwedish geologistAntevs,inthepagesofthePollenAnalysisCircular(oneofthefirstjournalsdevotedtopollenanalysis, and produced by Paul Sears in North America). Hyde and Williams chose palynology on the basis of the Greek wordspalunomeaning'tosprinkle'andpalemeaning'dust'(andthussimilartotheLatinwordpollen).Theyused thetermpalynologyforthestudyofexternal(morphological)featuresofmaturepollengrains. Palynology is an interdisciplinary science and is a branch of earth science (geology or geological science) and biological science (biology), particularly plant science (botany). Stratigraphical palynology is a branch of micropalaeontologyandpaleobotanywhichstudiesfossilpalynomorphsfromthePrecambriantotheHolocene.

History
Mohl(1835)classifiedthepollengrainsaccordingtotheirapertures,aperturemembranesetc.CarlJuliusFritzsche whosepaperUeberdenPollenpublishedin1837contributedtothestudyofpollengrainswithillustrationsand accuratedescriptions.HugoFischer(1890)describedthedetailsofexineinanumberofplants.Themoderneraof pollen analysis and the subsequent development of palynology, starts with Lennart von Posts lecture on Quaternary deposits in 1916. Roger P. Wodehouses classic Pollen Grains was published in 1935. Erdtmans Pollen Morphology and Plant Taxonomy published in 1952 is a monumental work in palynology. Text Book of PollenAnalysisofFaegriandIvesen(1964)isanotherimportantworkonprinciplesofpollenanalysis.Theadvent ofelectronmicroscopyputnewimpetustothestudyofthefinerdetailsoftheentirepollengrainsandspores.

Branches
PalynologycanbedividedintoBasic(Fundamental)andAppliedPalynology.BasicPalynologydealswithpollenand spore morphology, the physical and chemical properties of sporoderm as well as the correlation between palynologyandcytology.Italsoincludestheoreticalaspectsofappliedpalynology. AppliedPalynologydealswithappliedaspectsofthepalynology.Itincludes Palynotaxonomythestudyofpollenmorphologywithrelationtotheplanttaxonomy. GeoorPalaeopalynologythestudyoffossilpollengrainsorspores Melittopalynologythestudyofpollengrainsfoundinhoney Aeropalynologythestudyoffrequencyanddistributionofsporesintheair Iatropalynologythestudyofmedicalaspectsofpalynology,speciallytheallergies Pharmacopalynologythestudyofrolesofpollengrainsandsporesindrugs Copropalynologythestudyofpollengrainsandsporesinexcrements Forensicpalynologyapplicationofpollengrainsasanaidincriminologyandcrimedetection.

Applications
Importanceofpollenintheimprovementofplantswasrecognizedasearlyas717BCbyAssyrianartisans.Ancient Indian classics contain some reference about the refreshing pollen bath of Indian laburnum (Cassia fistula). The cosmeticchestsofEgyptianqueensarebeliegedtohavecontainedpollenpowderforfacialcare.Pollencultwas developed by American Indians who considered pollens of cattail (Typha) and corn as an emblem of peace, happinessandprosperity. Pollensarealsousedashumanfoodbysomepeople.AmericanIndiansusepollensofTyphaforpreparingpollen souporgruel.FewtribesofNewZealand,IndiaandPakistanuseTyphapollensasfoodbymixingwithwaterto formcakesandbaked.Themedicinalimportanceofpollenandsporeswaslongrecognized.Mentionmaybemade of Lycopodium spores used by the ancient Arabian physicians to cure stomach disorders, spores of Adiantum Philippines used in leprosy and skin diseases, pollens of Typha laxmanii used as astringent, Cycas circinalis as narcotic. Bee collected pollens mixed with coffee, milk, honey, potatoes and chocolates are prescribed to cure manydisordersinchildren. Significance of pollen in agriculture has been recognized in recent times. Selection of pollen with the desired potentialfromdesiredindividualsofanyonecroppopulationisconsideredasanessentialprerequisiteforplant breeding. Storage of pollen is as important as storage of seeds, in maintaining plant genetic resources. In many researchinstitutes,thehaploidplantsareusedforthemaintenanceofgeneticstock.ItisclaimedbyChineseand Japanesescientsthathaploidplantsofrice(Oryzasativa)developedfrompollenshavebeentransferredfromtest tubestofieldconditions.Insomecountries,storedpollensaredisseminatedovercropfieldsbymeansofsprayers drawn by tractors, or by breaking Pollen bombs or by hand, on days with moderately high winds. Successful pollinationensuresabundantfruitsuitandincreasedseedproduction. Pollensareidealmaterialforexperimentallycontrollingthegeneticbehaviorofplants,tproducemoreandmore usefulhybrids.Pollenirradiationisusedtomeettheproblemsofintraspecificorinterspecificincompatibility,by inducingdesiredmutationalchangesinthepollengenome.Inductionofmalesterilitybecomesaboonincrossing selfpollinatedplantslikegrapes. Aeropalynological studies of aeropollens have new vista in controlling the weed and has been practiced successfullyinEngland. Study of honey pollens provides information about the plants from which the nectar and pollens have been gathered.Suchstudiescandetectadulterationinhoney.SharmaandNair(1965)statedthatifhoneycontains50 percent of more pollens of one plant, it should be called Unifloral honey otherwise, it should be referred to as Multifloralhoney. Palynologyisusedforadiverserangeofapplications,relatedtomanyscientificdisciplines: Biostratigraphy and geochronology. Geologists use palynological studies in biostratigraphy to correlate strataanddeterminetherelativeageofagivenbed,horizon,formationorstratigraphicalsequence. Palaeoecology and climate change. Palynology can be used to reconstruct past vegetation (land plants) and marine and freshwater phytoplankton communities, and so infer past environmental (palaeoenvironmental)andpalaeoclimaticconditions. Organic palynofacies studies, which examine the preservation of the particulate organic matter and palynomorphs provides information on the depositional environment of sediments and depositional palaeoenvironmentsofsedimentaryrocks.

Geothermal alteration studies examine the colour of palynomorphs extracted from rocks to give the thermal alteration and maturation of sedimentary sequences, which provides estimates of maximum palaeotemperatures. Limnology studies. Freshwater palynomorphs and animal and plant fragments, including the prasinophytes and desmids (green algae) can be used to study past lake levels and long term climate change. Taxonomyandevolutionarystudies. Forensicpalynologythestudyofpollenandotherpalynomorphsforevidenceatacrimescene. Allergy studies. Studies of the geographic distribution and seasonal production of pollen, can help sufferersofallergiessuchashayfever. Melissopalynologythestudyofpollenandsporesfoundinhoney. ArchaeologicalPalynologyexamineshumanusesofplantsinthepast.Thiscanhelpdetermineseasonality of site occupation, presence or absence of agricultural practices or products and plantrelated activity areaswithinanarchaeologicalcontext.BonfireShelterisonesuchexampleofthisapplication.

Because the distribution of acritarchs, chitinozoans, dinoflagellate cysts, pollen and spores provides evidence of stratigraphical correlation through biostratigraphy and palaeoenvironmental reconstruction, one common and lucrativeapplicationofpalynologyisinoilandgasexploration. Palynologyalsoallowsscientiststoinfertheclimaticconditionsfromthevegetationpresentinanareathousands ormillionsofyearsago.Thisisafundamentalpartofresearchintoclimatechange.

MethodsofStudy
For the study of their external morphology, pollen slides are prepared by acetolysis as suggested by Erdtman (1952). Palynomorphsarebroadlydefinedasorganicwalledmicrofossilsbetween5and500micrometresinsize.Theyare extracted from rocks and sediments both physically, by wet sieving, often after ultrasonic treatment, and chemically, by using chemical digestion to remove the nonorganic fraction. For example, palynomorphs may be extracted using hydrochloric acid (HCl) to digest carbonate minerals, and hydrofluoric acid (HF) to digest silicate mineralsinsuitablefumecupboardsinspecialistlaboratories. Samples are then mounted on microscope slides and examined using light microscopy or scanning electron microscopy. Once the pollen grains have been identified they can be plotted on a pollen diagram which is then used for interpretation. Pollen diagrams are useful in giving evidence of past human activity (anthropogenic impact),vegetationhistoryandclimatichistory. Palynology uses many techniques from other related fields such as geology, botany, paleontology, archaeology, pedology,andgeography.

PollenMorphology
Pollenmorphologydealsmainlywiththewallsofpollengrainsandsporesandnotwiththeirlivingcontents. Thepollenwallhasaverycomplexstructurethatisreflectiveofthatspecificspecies'functionaladaptations.The exine is the outer layer of a living pollen grain. It is composed of sporopollenin, with small quantities of

polysaccharides.Sporopolleninisverychemicallystableanditisresistanttotoalmostallkindsofenvironmental damage. It is equipped with apertures. It it divided into two sub layers: the outermost sexine and the unsculpturedunderlyingnexine.Thesexinehassurfacesthataresculpturedinelaborateways,withreticulately arrangedperforations.Thesegivetheexineandamorphousorgranularappearance.Theinnerlayerofaliving pollengrainiscalledtheintine.Itiscomposedofcelluloseandisverysimilarinconstructiontoordinaryplantcell walls.Alayercalledtheendexineseparatesthesexineandintine.Theendexinehasalaminatedappearance. Pollen grains are generally classified according to their physical appearance. There are three criteria of classification:1)thenumberandpositionoftheapertures;2)theshapeofthepollengrainasawhole;and3)the fineelaboratestructureonthesexine.Aperturesareanymissingpartsoftheexine,whichareindependentofthe exinepattern.Aperturesarebigandtheycutacrossthefinestructurepatternonthesurfaceofthepollengrain. There are two types of apertures: pori or pores are mostly isodiametric apertures, although the can be slightly elongatedwithroundedends;colpiorfurrowsarelongandboatshapedwithpointedends.Colpiarethoughtto bemoreprimitive.Inlivingpollengrainstheseaperturesarenotactuallyopen.Instead,averythinlayerofexine coversthem.Grainswithporiarecalledporate;thosewithculpiarecalledculpate;andthosewithbothporiand culpi arecalled colporate. If their apertures are arranged equidistantly around the equator of thepollen grains theyareassignedtheprefixzono;iftheyarescatteredalloverthesurfaceofthepollengraintheyareassigned the prefix panto. The number of apertures is also indicated by prefixes: mono for one aperture; di for two apertures;triforthreeapertures;andsoon. Theshapeofapollengrainreferstotheshapeoftheiroutlineinpolarandequatorialviews.Theshapeofagrain cansometimesbeusefulinidentifyingofpollenspecies,butnotusually. Thesculpturereferstothefinestructureandpatternofthesexine.Itiscomposedofsmallradiallydirectedrods. If these rods support something (such as a plate or a small knob) they are called columnellae; if they do not supportanythingtheyarecalledbacula.Theshapeoftherodscanfurtherclassifythem.Iftheyareclubshaped they are called clavae; if they are sharply pointed they are called echinae; if they have swollen heads they are calledpila;andiftheyareshortandglobulartheyarecalledgemmae.Therearemanymoreclassificationsforthe shapeofrodsonthesurfaceofthesexine,butthesefourarethemostcommon. Ingeneralcharactersofpollensarestudiedunderthedivisionspollenunits,polarity,symmetry,apertures,size andshape,exinestratificationandexinepalynogram. PollenUnits Single pollen grains and spores, which when mature, do not remain united with other polleng rains and spores whicharereferredtoasmonads.Ifpollengrainsareunitedinpairs,theyarecalleddyas(Scheuchzeriapalustris, podostemonaceae).Atetradconsistsoffourpollengrains.Anoctadofeight,apolyadofmorethaneightpollen grainsarealsofound. Tetradscanbecategorizedasfollows Tetrahedraloneofpollengraisnlyingcentrallyovertheotherthree(Drimys) Tetragonal(Isobilateral)allpollengrainsinoneplanebluntlyquadrangular(Monocotyledons) Rhomboidal(Anonamuricata) Decussate(Cross)pollengrainspairwiseatrightanglestoeachother(Magnoliarandiflora)

Tshaped the first division of the PMC is ransverse and a dyadcell isformed, either the upper orthe lowerdyadcellundergiesverticalorlongitudinaltoyieldinvertedTshapedconfiguration(Aristolochia) Linear the first division is always transverse and gives rise to dyad cells and second division also transversetogivelineararrangement(Mimosapudica).

Insedges(cyperaceae)monadliketetradsareformedwithoutdividingwallsbetweenthefourcomponents.One nucleusisnormallydeveloped,whereastheremainingthreeareobliterated.Suchtypeofdegradedtetradiscalled pseudomonad. InMimosaceaeeachofthetradcellsdividesonceortwiceandthesearisesamassulaeofeightto64cellswhich remaintogetheraftermaturity,(butnottheentireproductofonetheca).Theyconstituteasmallunitsandalso calledpolyads.

Inpolliniumaunitofallgrainsproducedbyonetheca(loculeinsidetheanther)aggregatetogethertoformone singleunit.Anentirepollenmassofathecalchamberformthepollinium.Thepolliniumapparatusisthefunctional unitofacorpusculumwithitstwoattachedarmsandpollinia(Calotropis,Orchidaceae) Polarity All pollen grains have a tetrad stage during development. It is at this stage that polarity is determined. At the tetradstage,theendofapollendirectedtowardsthecentreoftetradiscalledproximalpolewhileoppositeisthe distalpole.Thedisalpolefacesperipheralside.Thepolaraxisisalwaysperpendicularwiththedistalpoleatthe apeandtheproximalpoleatthebase.Thecentreofeachpatiscalledpoleandtheimaginarystraightlinethrough thepolesiscalledthepolaraxisandeachplanethroughthisaxis isa polarplane.Theborderlinebetweenthe distantpartandtheproximalpartofthepolengrainistheequatorandtheplanethroughtheequatoriscalledthe equatorialplane.

 Pollengrainsmaybe Apolarpollengrainswithoutdistinctpolarity,forexample,spheroidalgrains,thatiswithoutdiscernible polesoncethegrainshavebeenseparatedfromthepollentetrads Polarpollengrainswithrecognizablepolesevenafterthegrainshavebeenseparatedfromthepollen tetrad.

Inisopolargrainsthedistalandproximalpoleslookalike.Inheteropolarthetwofacesareessentiallydifferent. Grainswhichhavemoreorlesscurvedequatorialplaneandpossessonlyslightdifferencesbetweenthedistaland proximalfaces(oneconvesandotherconcave)maybetermedparaisopolar(subisopolar).Inlineartetradthereis nopolarityintheusualsenseoftheterm. Symmetry It is the quality inherent in a body which is capable of division into similar or equal halves. Spores may be symmetrybeingbilateralorradial.Inradiosymmetricsporestheshapeissuchthatanyplaneincludingthepolar axisthatpossesthroughwillproduceidenticalhalves.Bilateralsporeshavetwoverticalplanesofsymmetry.Some spores are asymmetric (with no symmetry at all). Some spores possess fixed shape (fixiform) while some are withoutafixedshape(nonfixiform). Shape AccordingtoErdtman,radiallysymmetricalpollengrainsareshapedlikearotationellipsoid,withthepolaraxisas theaxisofrotation.Theshapeiscalculatedbytakingmeasurementsofpolaraxis(P)andthemaximumbreadth(E) intheequatorialviewofthegrainandapplyingtheformulaP/EX100.Thevaleobtainedgivestheshape. Theshapeclassesareasfollows ShapeClasses Peroblate Oblate Subspheroidal Suboblate Oblatespheroidal Prolatespheroidal Subprolate Prolate Perprolate P/EX100 <50 50 75 75 133 75 88 88 100 100 114 114 133 133 200 >200

Confervoid grains (filiform, filamentous) are described as long simple, one walled threads arranged in parallel massesequallyingtheantherinlengthandtendingtobecoeverytangledwhendisturbed.(Zosterasp)