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Age States of Plants of Various Growth Forms: A Review Author(s): L. E. Gatsuk, O. V. Smirnova, L. I. Vorontzova, L. B. Zaugolnova and L. A. Zhukova Source: Journal of Ecology, Vol. 68, No. 2 (Jul., 1980), pp. 675-696 Published by: British Ecological Society Stable URL: http://www.jstor.org/stable/2259429 . Accessed: 19/09/2013 20:12 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. . British Ecological Society is collaborating with JSTOR to digitize, preserve and extend access to Journal of Ecology. http://www.jstor.org This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions Journalof Ecology(1980), 68, 675-696 AGE STATES OF PLANTS OF VARIOUS GROWTH FORMS: A REVIEW L. E. GATSUK, 0. V. SMIRNOVA, L. I. VORONTZOVA, L. B. ZAUGOLNOVA AND L. A. ZTIUKOVA of Botany, BiologicalProblemLaboratory,Department 129243Moscow,USSR LeninState Teachers'TrainingInstitute, SUMMARY plant specieshas showncertaincon(1) Analysisof morethan one-hundred Theseare calledseed, biologicalage statesin theirontogeny. sistently-recognizable (young,matureand old), seedling,juvenile,immature,virginile,reproductive subsenileand senile. of quanticombination bya particular (2) Each age statemaybe characterized tativeand qualitativefeatures.Qualitativefeaturesused to definethevariousage thetypeof growth,thepatternof branching statesare: themannerof nutrition, leafform,thepresenceofa particular typeofshoot, oftherootand shootsystems, theabilityto reproduceby seeds,thebalancebetweenlivingand dead structures, and fully-formed structures. and thebalancebetweenactively-growing duringontogeny,and changeuninterruptedly (3) Quantitativecharacteristics as a rulefollowa unimodalcurve. of a varietyof growthformshave been (4) Age statesof speciesrepresentative low semiand described.These includetrees,shrubs,semi-shrubs, distinguished plants,and thefollowingcategoriesof perennial shrubs,firm-and loose-tussock stoloniferous, bulbous,tuberroot-suckering, herbs: long-and short-rhizomed, bulbous,tuberousand tap-rooted. (5) On the basis of theseage-statestudies,threemain typesof ontogenyin changes features architectural usingas characterizing plantsare defined, polycarpic of theindividualplant,theformand timingof break-upof theindividualplant, and theextentof rejuvenation. INTRODUCTION Living organismsfrombirthto death undergo sequentialphases which are usually of an indivicharacterized bychronologicalage. This sequenceofphasesof development anatomical,physiologicaland dual organismis called ontogeny.Variousmorphological, biochemicalchangestake place in the course of ontogeny,so that a givenindividual not plant or animal at a particularmomentof its developmentmay be characterized, onlybyitscalendarage, butalso bybiologicalcriteriawhichindicateitsstateofdevelophas beenvariouslycalled'biologicalage' (Robbins 1957; ment.This secondcharacteristic de Muckadell 1959; Grubb 1977), Levin 1966), 'physiologicalage' (Schaffalitzky 'ontogenicage' (Passecker1977) or 'age state' (Uranov 1975). As a measureof biological age in plants it is possible to defineage stateswhich representsuccessivestages of theirontogeny.The indicatorsof age statesare mostly morphologicalchanges,whichcorrelatewithanatomical,physiologicaland biochemical Publications ?1980 BlackwellScientific 0022-0477/80/0700-0675$02.00 675 This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions 676 Age statesofplants changes(Krenke1940;Sinnott1960;Gupalo 1969).Each succeedingstageis characterized whichwerelackingin earlierstagesof development, bytheappearanceof newstructures and by the loss of previousones (Uranov 1975). but since The averagedurationof age statesof each plantspeciesis fixedgenetically, individualsmay in factreach a conditionsmay varygreatly,different environmental dates. In a spruceforest,forexample,theremaybe some certainage stateat different 70-yr-old treesthathave neverproducedanyseedswhileothersofthesamecalendarage are activelyreproducing. considerthe calendarage of plants(Rabotnov Most researcheson plantdemography 1950; seeHarper& White(1974) fora review).However,we believethattherecognition and than calendarage in analysingthe structure of age stateshas greatersignificance have age statesdo notnecessarily dynamicsofpopulations.Individualplantsof different roles in the lifeof a coenosimilarecologicalproperties,and theymay play different (Rabotnov 1950, 1974). The renewaland ontogenetic populationor a plantcommunity of plantsresultsin a complexutilizationof habitatresourcesbothin space development to the coexistenceof many species in and in time,and may contributesignificantly (Watt1947; Rabotnov1969; Kershaw1973; Uranov 1975; diversified plantcommunities is determined morebyits Grubb 1977).Thus therole of a givenplantin thecommunity age statethanby thenumberof yearslived; henceit is moreimportantto knowits age individuals state.Similarage statesmaybe attainedby different species,and bydifferent ofthesamespecies,at different calendarages,and a comparativemeasureoftheirrolein a community on thebasis of calendarage has onlyverylimitedbiologicalmeaning.The definitionof calendarage of manyherbsand woody plants,moreover,is practically impossible,whereasa classificationby age statesis quite feasible.The simultaneous of bothcalendarand biologicalage is mostuseful,ifit is possible. definition DEFINITION OF AGE STATES The ontogenyof plants may be dividedinto periods or stagesbased on the rise and of thereproductive functionand on certainjuvenileand adultcharacteristics: extinction on this basis pre-reproductive (juvenile), reproductive(adult) and post-reproductive However,more detaileddivisionof the ontogenetic (senile) periodsare distinguished. sequence may be achievedusing the process of formationand developmentof plant theirchangesand death,and thecorrelationoftheseprocessesin an organism. structures, of an individualand its outcome (e.g. vegetative The onsetof physicaldisintegration particularage states. propagation)mayalso be usefulin defining botanical ofage stateswellknownin theRussian-language Therearetwoclassifications fortrees(Schitt1958),the otherinitiallyfor literature:one was developedspecifically perennialherbaceousplants(Rabotnov 1950) but has since been more widelyapplied (Uranov 1975; Smirnovaet al. 1976; Uranov et al. 1977). The currentsystemis summarizedin Table 1. theontogenyofabout 100 UndertheguidanceofA. A. Uranovand I. G. Serebrjakov, These includea plant(listedin AppendixI) have been investigated. speciesof flowering widerangeoflifeforms(fromtreesto annuals),withvariedecology,geographicaldistrihas beenfoundapplicableto butionand systematic status,and an age stateclassification all thespeciesstudied. To studytheontogenyof a particularspeciesand to describeitsage states,theplants collectedin such a way thatthewholeindividualplantis preservedintact. are carefully This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions 677 L. E. GATSUK et al. TABLE1. Ontogenetic periodsand age statesofplants Ontogenetic period Agestate Symbol I. Latent (1) Seed se II. Pre-reproductive*(2) Seedling pI (3) Juvenile j (4) Immature im (5) Virginile v III. Reproductive (6) Young gl (7) (8) IV. Post-reproductive (9) (10) Mature Old Subsenile Senile 92 g3 ss s * A more exact translation of the Russian terms wouldbe pre-generative, generative andpost-generative Large trees are describedand measureddirectlyin the field.For each individuala morphologicallyand physicallycontinuous specimen,isolated from the others,is selected and analysed. Aftercareful morphologicaldescriptionthe individualsare arrayedin an ontogenicseries,fromseedlingsto senileplants.The ontogeneticseriesis thenpartitioned intoage states,dependingon thepecularitiesofplantstructure. Between tenand thirty normally-developed individualsare used to describeeach age state. In thispaper descriptionsof age statesof fourplantspeciesof different lifeformare given: a tree,FraxinusexcelsiorL. (by L. B. Zaugolnova); a tap-rootedsemi-shrub, Anabasis aphyllaL. (by L. E. Gatsuk and L. I. Vorontzova); a firmtussock grass, DeschampsiacespitosaBeauv. (by L. A. Zhukova); and a long-rhizomed perennialherb, Aegopodium podagrariaL. (by 0. V. Smirnova). DESCRIPTIONS OF AGE STATES Fraxinusexcelsior zones of Europeand in Fraxinusexcelsioris widespreadin theforestand forest-steppe themountainsof Asia Minor.Usuallyit occursas a subordinatespeciesof thetreelayer but in some standsit maybe dominant(Boysen-Jensen 1932). Eight age states were distinguished by Zaugolnova (1968) in the ontogenyof ash growingin thebroad-leavedforestsof theEuropeanpartof theUSSR (Fig. 1(a)). (1) Fruit.A single-seedednutlet,usuallygerminating onlyin the year subsequentto shedding(Wardle1961). (2) Seedling(Fig. l(a), pl). Appearsin late April,and by the end of thefirstgrowing season undera forestcanopy has a hypocotyl,an epicotyl,two cotyledons,a pair of simpleleaves and an unbranchedtap-root. (3) Juvenile plant-calendar age 2-15 yr,height0.1-0.4 m (Fig. 1(a), j). Differsfrom the seedlingin theabsenceof cotyledonsand thepresenceof compoundleaveswith3-7 leafletsand an unbranchedmonopodialshoot; theroot systemis composedof a vertical tap-rootand horizontallateralroots.The juvenileplanthas a highshade-tolerance. (4) Immature plant-calendar age 10-20 (maximum27) yr,height0.5-3.5 m (Fig. 1(a), im). In thisage state,matureleaves with9-13 leafletsare present,and a monopodial branchesdevelops,thoughnot forminga shoot systemwith second-and third-order witha tap-rootwhichis shorterthanthe definitecrown;the root systemis superficial, increasessharply,and immatureplants second-order lateralroots.The light-requirement are foundonlyin sunlitareas of a forest. This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions 678 Age statesofplants Cd 0 C.) II.4 0I N - - ~~~~0 (0 ~~~~~~~~~~;IN 2 $I~~~~~~ti~~~7>~ 0 N- - 0 C4 C~~~~~~~4- E 0 0 LO 0 ~~ LO 00 0~~ 0 0) N E L L A~~~~~~~~~~~? L~~~~~~~ U~~~~~~ 0N 0 r~~~~~~~~ ~0 ~0 ~-O~~~~~~~~~~~ ~ ~ ~ N _ - ~ 0~~~~ _~ ~ .. ~ ~ ~ C This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions C L. E. GATSUKet al. 679 plant-calendar age 20-30 yr,height4-10 m (Fig. 1(a), v). A typicaltree(5) Virginile trunksurmountedby a branchedcrown,is present;the form,with a clearly-defined withthe tap-root trunkis coveredwithperidermonly; the root systemis superficial, persisting.Virginileplants neitherflowernor bear fruit,but show maximumannual leader-shootgrowth. plant-calendar age 30-50 yr,height10-15 m (Fig. 1(a), gl). (6) Youngreproductive fastigiatecrownwiththe maximumwidthin its lowerpartis present; A well-branched ratherthinand smoothbarkdevelopson thetrunk;therootsystemis varied,depending begin. on soil conditions.Floweringand fruiting plant-calendar age 50-90 yr,height17-25m (Fig. 1(a), g2). (7) Maturereproductive crown,up to 8 m in diameter,is present,withmaximumwidth A largerounded-rhombic in themiddleor upperpart; thetrunkis coveredwiththickbarkwithsmallcracksin it. Severalprocessesindicativeof ageingoccur: reducedannual shoot elongation,delayed leader-shootelongation,die-backof weak lateralbranches,dyingof large branchesin of thebranchingzone to theapical partsof shoots. thecrown,and thetransference plant-calendar age 100 yr or more,height25-30 m (Fig. l(a), (8) Old reproductive withmaximumwidthin its upperpart; thebark on g3). A moreopen crownis present, the trunkis deeply fissuredas a resultof dead tissue accumulation.Annual shoot elongationdiminishessharply,and the death of axillaryand terminalbuds reduces branching;thosebranchesthatdo formare usuallysmalland twisted.Senescencein the root systemis apparentas a decreasein thevigourof branching,and thedyingof root tipsand woodyroots. A post-reproductive period(subsenileand senileage states)does not occurin ash; old treesfruituntiltheirdeath (150-220 yr). The ontogenyof severalspeciesoftimberand fruittreeshas beenstudied(AppendixI). The followingcommonpatternscan be recognized;(a) a branchedcrownis formed period(20-60 pre-reproductive beforeflowering begins,whichoccursonlyaftera lengthy withage; (c) to rounded from crown-form fastigiate changes yrforforesttrees);(b) the theonset coincides with and in trunk diameter maximumincreasein annualshootlength of is with evident combined signs ageing (d) maximumfruiting of flowering and fruiting; annual shoot elongation,reductionof (decrease of terminalleader-shootgrowthand of die-back and branches); (e) ageing is expressedby currentannual trunkgrowth, progressivedeath of crownbranches,twistingof branches,reductionof fruitingand rootingcapacity,and decreaseof annual shootelongationand cambialgrowth. Anabasis aphylla and desertin Kazakhstan,thelowerpartsofthe Anabasisaphyllaoccursin semi-desert RiversVolga and Don, and in Kashgaria and Jungaria.The plant is a dominantor of manylowlandand foothilldesertcommunities anabasetum). (Artemisieta co-dominant withlong yearlyshootswhichappear Anabasisaphyllais a halo-xerophytic semi-shrub, leaflessbecausetheleafblades are reduced.The steminternodehas a leafcylinder('leaf sheath')attachedto it (Leysle 1949). The upperpartof a yearlyshootis deciduous,and and succulent at theend ofthevegetativeseasonit dies off;thisis branched,assimilating (Fig. 2, (Fig. 2, I). The lowerpartof a yearlyshootmaybecomeperennialand persistent II), and build a skeletalbasis to the plant. Persistentparts of some sequentialyearly shoots('podia') forma sympodial'skeletalbranch'(Fig. 1(b), g2, 7), whichlivesfrom3 ofwoodytissue. to 5 years.Persistent partsof theyearlyshootsconsistalmostentirely This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions Agestatesofplants 680 * a [II 10 2 6 4 2 1p0 individual aphylla. (g2) ofAnabasis shootofa mature reproductive FIG. 2. A stylized annualshoots, part;1 = deciduous I = deciduous shoot,II = persistent partofyearly 2 = flowers 3 = nodes. and fruits, reach75 cm in heightand Largerspecimensof Anabasisat theirpeak of development The tap root 130 cm in diameter.The plant has no abilityto propagatevegetatively. increases and thereare no diameter its basal every year, has a life-span, long system of the plant's absolute age by morphoadventitiousroots. The correctdetermination This is due to repeatedshootbranching logicaland microscopicmethodsis verydifficult. die-back of yearlyshootsto the base, shoot formation duringeveryvegetativeseason, on and a buds the complicatedpolycambialanatomicalstructure. fromadventitious root, that data Indirect relatively largespecimensin naturalconditionslivefor20-25 suggest Nine age-statescan, however,be recognizedwithoutdifficulty. years. non-endospermous. (1) Fruit.Large, single-seeded, (2) Seedling.Emergesin late Aprilor earlyMay. The seedlinghas onlya tap-rootand a main shoot; both cotyledonsare stillgreen. plant(Fig. 1(b), Fig. 3; j). Cotyledonsare dry;twoto fourgreenjuvenile (3) Juvenile blades are present;subsequentlyleaves of transitionaltype leaves withclearly-visible This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions 681 L. E. GATSUK et al. cm 4- 3 3- 2 2 0 Im plants of Anabasisaphylla;j = juvenile(mid-May),im= FIG. 3. Pre-reproductive 2 = leavesofjuveniletype,3 = leavesof (mid-June);1 = dead cotyledons, immature type. transitional is stillrelatively simple,with withreducedbladesmaydevelop.The plantstructure shootandroot. unbranched smaller plant(Fig. 1(b),Fig. 3; im).Newleaveswithbladesnoticeably (4) Immature andat theendsoftheshootstheleafbladesare thanthoseofjuvenileleavesarepresent, plants reducedandtheleafsheathsarefusedto theshootaxisas in mature completely of feature and oftenalso thejuvenileleaves,die.A characteristic (v-s).Thecotyledons, lateralshootsarisebasitonicis theonsetofbranching: age-state plantsintheimmature buds. axillary above-ground allyfrom notonly is present, branching axillary plant(Fig. 1(b),v). Moreextensive (5) Virginile (deciduous) partoftheshootbutalso intheupperassimilating inthelower(persistent) part.Allleafbladesarereduced. symmetrical andrelatively plant(Fig.l(b),g,).Tallandcompact (6) Youngreproductive areborneon the and fruits and buds;flowers deadbranches evident in shape,without in thefirst mostlargeplantsflower deciduouspartoftheyearlyshoots.In cultivation, 'skeletalbranch'system(Fig. l(b), g2, 7) is A sympodial autumnaftergermination. (podia) or olderplants,and thelengthof itsannualincrements in 2-year-old present eachyear. increases This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions 682 Age statesofplants plant (Fig. l(b), g2). Numerousvigorousbranchedshoots, (7) Mature reproductive materialand strong'skeletalbranches',are present. witha highproportionof persistent Plants of this age state are characterizedby the greatestvegetativemass and seed At the same timeclear signsof ageingare and a largeskeletalframework. productivity, present:the lengthsof successivepodia in each sympodial'skeletalbranch'begin to decreaseeveryyear,and theinitialbasal clusterof shootsand thebasal partof thetaprootdie back partlyor wholly.However,not manydead partsoccur,and theplantas a wide and large. ruleremainssymmetrical, plant(Fig. 1(b), g3). The initialclusterof 'skeletalbranches'dies (8) Old reproductive buds at thebasal adventitious back wholly.Vigorousshootsdevelopfromunderground shootsgiveriseto part(i.e. nearestto thesoil surface)ofthetap-root;theseadventitious bush thusforminga flattened new 'skeletalbranches',whichusuallygrowhorizontally, withno shootsin thecentre.Some of these'skeletalbranches'thendie off,and thebush witha hollowor sectorof dead tissuein thebasal becomesone-sidedand asymmetrical, part of the tap-root.The lengthsof successivepodia of each 'skeletalbranch'steadily decrease,and yearlyshootsare muchshorterand thinnerthanthoseof matureplants. Thereare manydead organs.Reproductivecapacityis weakened. (9) Senileplant (Fig. 1(b), s). The plantno longerbears fruit,and the signsof senescenceare moreevident.'Skeletalbranches'arisefromthetap-rootfroma depthofup to bush.The basal partofthe 30 cm,butfromone side only,givingriseto an asymmetrical rootis oftenhollow,and althoughtheroot diameteris large,onlya partof the root is activelygrowing.Few new shootsare produced,and theseare short,thin,unbranched or weaklybranched,and oftenwhollydeciduous.Thereare fewor no viablebuds. Anabasisaphyllahas certainfeaturesof its ontogenywhichare also shownby some life(AppendixI). In all and herbswhichpreservethe tap-rootthroughout semi-shrubs theseplantsthe centralclusterof shootsand the core of the basal part of thetap-root dies offtowardstheend of ontogeny.As a resulttheabove-groundsystemmaybecome as in Artemisia compacta. dividedintoindependent plants(completeseniledisintegration), in a similarspatialposition Anabasisaphyllahas certainuniquefeatures:thereplacement of 'skeletalbranches'of shoot originby ones of root origin;the reductionof the leaf blade fromthevirginileto thesenilestates;and theabsenceof leaves ofjuveniletypein thesenilestate. Deschampsiacespitosa Deschampsiacespitosa,a tussockgrass,occurswidelyin wet and drygrasslandand alpinemeadowand in theforestsofWesternand EasternEurope,Asia, Africa,Australia In and Northand SouthAmerica.It is a polycarpicperennial,obligatelymycotrophic. plantsfromtheplainsand alpinemeadowsof theEuropeanpartoftheUSSR, Zhukova nineage states(Fig. 4(a)). (1961) distinguished withouta dormancyperiod. (1) Fruit.A smallcaryopsis,oftengerminating (2) Seedling-calendarage from2-3 weeksto severalmonths(Fig. 4(a), pl). Consists leaves,radicle,and 4-5 of a primaryshoot,withcoleoptileand 2-3 narrowassimilating remainsconnected The cm in each roots 3-5 seedling length. adventitious non-branching to thecaryopsis. (3) Juvenile-calendarage froma fewmonthsto 5-6 yr (Fig. 4(a), j). Consistsof a shootwith6-7 leaves,2-3 of whichmayhave primaryunbranchedor slightly-branched not as largeas thoseofmatureleaves;theadventitious died; theleafbladesare larger,but This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions L. E. GATSUK et a!. 683 tt II U)~~~~~~~~~~~U 0 o o ~~~~~~ U) 0 ~ ~~~~~~~~~~~Ir~- ;* 0 - P ..: o 1" - 0 S 0~~~~ 0 ~ tw II I. 0~ > e) 0 ~~~~~~~~~~~~~~~~~~~~~~~~~o:U This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions 00 0 684 Age statesofplants and reach a depthof 12-15cm. The connectionwiththe caryroots branchslightly, opsisis lost. age-stateof Deschampsiacespitosais practicallyimpossibleto distinThe immature features. guishbymorphological (Fig. 4(a), v). In the virginilestate,the typicalgrowthformof a firm(4) Virginile a roundedtussock,8-10 tussockgrassbecomesevident.As a resultof intensivetillering cm in diameter,is formed.An emptyzone devoid of vegetationappears around the of Deschampsiaon itsneighbours. influence tussock,perhapsas a resultofthephytotoxic Leaves typicalof matureplantsare formed,withblades 25-30 cm long and up to 2 cm wide. The root systemincreasesin width and depth (18-25 cm), because of newlyroots. adventitious formeed The durationof thevirginilestateis 2-3 yr,or up to 5 yrin unfavourableconditions. D. cespitosafirstflowerswhen6-7 yr In naturalcoenosesin favourableenvironments, periodoftussockgrassesare based on theproold. The subdivisionsofthereproductive portionsoflivingand dead partsin thetussock,theformofthetussock,thesizeand form each tussock,and the lengthof the 'pseudorhizome'. of the 'emptyzone' surrounding (Fig. 4(a), g,). The tussockis eitherround or oval in shape, (5) Youngreproductive withno dead parts.The emptyzone aroundthetussockis a wide ring.The root system continuesto increasein size each year.Reproductiveshootsconstituteonly3-4y/of all shoots.The plagiotropic(horizontal)partof theshootsis seldomevident. (Fig. 4(a), g2). Necroticpartsare formedin thecentreor at the (6) Maturereproductive of the rimof thetussockas so;ne shootsdie. This necrosisresultsin the disintegration 'primary'individual(whicharosefromseed)intoseparatepiecesor 'particules',separated by dead parts,but fora long timethe clone remainsin the formof a tussock.In the matureage-state,one or two of thelowerinternodesof manyshootselongateand form the zone of tilleringhigherand allowing so-called pseudorhizomes,thus transferring some parts of the tussockto maintainmorphologicaland physiologicalconnection. shoots Aroundthe tussockis a wide circularemptyzone. The numberof reproductive of any age state. increases,to c. 10% of all shoots,givingthehighestseed-productivity is maximal. The rateof lateralshoot-and root-formation (Fig. 4(a), g3). Plantsof maximumdiameter(0-3-1-0m), butwith (7) Old reproductive in theisolation morethanhalfthecentreofthetussockoccupiedbydead shoots,resulting of 2-20 particules,each of manyshoots.The clone of particulesremainscompact,and is stilltakenas theunitforpopulationcensus.The tussockis surroundedbyan emptyzone plants.Rate of formation by neighbouring of irregularform,whichis partlyovergrown of rootsand shootsis reduced,and some youngshootshave no roots at all. Thereare and 2-3 tillering shoots.All theseshootsform3-4 elongatedinternodes fewreproductive zones, so thattheyoungshootsare removedsome distancefromthedead centreof the tussock. The totaldurationof thereproductive periodis 15-30yr(Persikova1959). is shortened;as a result diminishes and shootlongevity (8) Subsenile.Tilleringintensity the numberof livingshootsin each particuledecreases,and the total area of the dead portionincreasesgreatlyand is invaded by otherspecies.Particulesare weakened,and the emptyzone around the tussockbecomesindistinct.Root formationand growthis shootsare formed.The majorityofrosetteshootshave rhizomes reduced.No generative formedby3-6 elongatedinternodes. (9) Senile.The tussockbreaksup, and onlytwo or threeparticulespersistamongthe dead parts. Tilleringis rare. Pseudorhizomesreach theirmaximumlength,as nearly This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions L. E. GATSUK et al. 685 all internodesare elongated.A few erect vegetativeshoots develop, bearingleaves of a juveniletype,withshortnarrowblades. The depth of the root systemdecreases sharply.The emptyzone aroundthetussockis indistinct. period is 15-25 yr. Thus the overall The total durationof the post-reproductive ontogenyof Deschampsiacespitosalastsfor25-60 yr. The ontogenyof severalfirmand loose-tussockgrasseshas beenstudied(AppendixI). passesthroughseveraldevelopmental Certaincommonfeaturesareapparent.The tuissock particulesand then stages,froman individualof seed originto a clone of closely-packed to a singleresidualparticule.Usually,shootdie-backstartsin thecentreof thetussock and spreadscentrifugally. grassesis The ontogenyof steppeand meadow firm-tussock typicallyof long duration,the reproductive(and in some cases post-reproductive) periods being especiallyprolonged.Loose-tussockgrasseshave a shorter(4-5-times) grasses,and typicallybegin to flowerafter pre-reproductive period than firm-tussock 1-3 yr. podagraria Aegopodiumn Aegopodium podagrariaL. occurs widelyin Europe, exceptthe extremenorthand south,and in Asia Minor and Siberia.It is a polycarpicperennialwithlong rhizomes. nine age statesin the ontogenyof A. podagraria. Smirnova(1967b, 1974) distinguished in theyearafterripening, in May or earlyJune. (1) The seed germinates for 1-2 (2) The seedling(Fig. 4(b), pl) has two cotyledons,whichphotosynthesize monthsbeforedying.The plantquicklydevelopsa primaryrosetteshoot,withone (or sometimestwo) 3-lobed leaves. During the firstyear the main root grows5-7 cm in length,and startsdrawingtherosetteshootdownintothesoil. (3) Thejuvenileage-state(Fig. 4(b), j) lastsfor2-3 yrin naturalhabitats.The cotyleleaves develop.The absoluteage of dons are lost,and two or three3-lobedassimilating youngindividuals(beforethe startof vegetativepropagation)may be determinedby countingthe numberof leaf-scarson the initial rosetteshoot, and dividingby the numberof leaves developingeach year (Shik 1953; Rabotnov 1960). Lateral roots develop,and adventitiousrootsemergeon thehypocotyl. (4) In the immatureage state (Fig. 4(b), im), 5-7-lobedleaves, transitionalin form betweenjuvenileand matureleaves,develop.The mainrootformsmanylateralsand the numberof adventitiousrootsis increased.At the age of 5-6 yrimmatureplantsform plagiotropicundergroundrhizomesfromaxillarybuds on the primaryrosetteshoot. These rhizomeswithelongatedinternodesand scale-shapedleaves are called runners (Fig. 4(b), 7). The lengthof the firstrunnersis 5-10 cm. At the end of autumneach and a new rosetteshoot developsfromthe runnerturnserect(becomes orthotropic), terminalbud; thisis calleda 'partialshoot'(Fig. 4(b), 8). The plagiotropicsegmentofthe rhizome'( Fig. 4(b), 9). runnerafterdevelopinga partialshootis calleda 'communicating consist of a rosette shoot and some partial primary (5) Virginile plants(Fig. 4(b), v) shoots joined to the primaryone by communicatingrhizomes.Each partial shoot develops an adventitiousroot system,includingthickstorageand thinfeedingroots rootsare formedat thenodes of communi(Fig. 4(b), 11). In addition,thinadventitious of relativelyautonomouspartialshoots, now consists individual rhizomes. The cating each withits own root systemand separatedfromthe primaryshoot by a distanceof to as polycentric. 20-30 cm. This morecomplexplantis referred Virginileplantsconsist is 200-300cm. The of total rhizomes 10-20 and the plagiotropic of partialshoots, length is in natural habitats. of 5-7 absoluteage virginileplants yr This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions 686 Age statesofplants period,or somewhatearlier,vegetativeseparation At the onset of the reproductive begins;as the oldestpartsof the plagiotropicrhizomesdecay,subdivisioninto two or several'filial'plantsoccurs.Filial plantsusuallyconsistof severalpartialshootsjoined fromthematernalplantbytheabsence byplagiotropicrhizomes,butas a ruletheydiffer is a systemof partialshoots. of a primaryrosetteshoot.At thisstagetheplantstructure The partial shoot has its own age, whichmay be called 'provisory'in contrastto 'absolute' age. We considerabsoluteage as thattimefromthe momentof germination of a seed untilthemomentof observation.The provisoryage (or 'properage'-Krenke 1940) is definedas the timefromthe inceptionof the oldestpreservedpart of a plant betweenabsoluteand provisoryage is untilthemomentof observation.The distinction and this confusioncan lead to incorrect not always clearlymade by investigators, estimatesofplantlongevity. bysystemsofpartialshoots, plants(gl-g3) are mostlyrepresented (6)-(8) Reproductive indivi'monocentric'generative Rarely,followingdisturbance, and are thuspolycentric. duals are formed,consistingof onlyone partialshoot; however,newpartialshootsthen again. arisefromrunners,and the plantbecomespolycentric by the proAge statesof the generativeperiod of Aegopodiumcan be distinguished and of old ones whichhave ceased their structures portionsof youngactively-growing are thoserunnersand partialshootswhichare capable of growth.The youngstructures formingnew runnersand adventitiousroots each year; theirprovisoryage is 1-5 yr. They may be called 'prospectivepartial shoots'. Old structuresare communicating rhizomesand partialshootsincapableof formingrunnersand adventitiousroots; these shoots have a small assimilatingsurfaceor are completelydevoid of green leaves partial ('stumplets');theirprovisoryage is morethan 6 yr.These are 'non-prospective shoots'. plantcan be definedon thebasis of two The age-stateof theindividualreproductive theproportionsof young coefficients, DI and a,,. The firstindexof age-state(a,) reflects the secondindex(a,,) tufts); shoots (or partial old (non-prospective) and (prospective) and communicating (runners) rhizomes growing actively of reflectsthe proportions rhizomes. p-P 'p+P Pi_p1P1 _ Pi +Pli where P = numberof youngpartialshoots(or tufts)of an individual; p = numberof old partialshoots(or tufts); P1 = numberof runners; rhizomes. Pi = numberof communicating for DI shows the generalageing of an individual,while 9II definesthe potentiality seen,forexample,in theperiodicformationof newrunners. partialrejuvenescence, plants (Fig. 4(b), g,) are composed mainlyof youngpartial (6) Youngreproductive shootsand runners(-1.0 < cI < - 0.5). The numberof partialshootsis small(mean 3.87),and thetotallengthofrhizomesaverages135cm (Smirnova1967b).The provisory age ofindividualsis 5-6 yr. (Fig. 4(b), g2) have thegreatestnumberof generaindividuals (7) Maturereproductive tive shootsand rhizomes;the mean numberof partialshootsis 4.5. Leaf size and the increases.Total lengthofrhizomesis on average155cm(longest degreeofleaf-dissection This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions L. E. GATSUKet al. 687 600 cm). Mature plants show vigorousgrowthof all organs,but theybegin to show signsofsenescenceas thenumberofyoungpartialshootsdecreases(-0.4 < a, 0-05). rhizomesalso decreases. The numberof actively-growing plants (Fig. 4(b), g3) show signs of ageing more clearly.The (8) Old reproductive numberofpartialshootsis reducedto 2.8 on average,and old shootsincreaseproportionately(-0.4 < a, < + 1.0). The numberof runnersdecreasessharply,as does the total period,plantslose the lengthof rhizomes(mean 84 cm). At theend of thereproductive (that is, they do notJformrunners),but vegetative capacityto expand vegetatively continues. fragmentation (9) Subsenileplants(Fig. 4(b), ss.). The numberof partialshootsis verysmall(1.5 on ofimmature average).The numberof leavesand theirdegreeofdissectionis reminiscent plants.These individualsformneitherrunnersnor new adventitiousroots. by singlepartialshootswithleaves of (10) Senileplants(Fig. 4(b), s) are represented juveniletype.From theappearanceof theaerial shootthesesamplesmightbe mistaken forjuvenileplants,but thepresenceof theperennialrhizomeindicatestheirsenility. to thepost-reproductive period, fromthereproductive Thus as a plantis transformed increases,leadingto the appearanceof secondarymonoso its degreeof disintegration occurin several certainindividualstructures centricindividuals.In theoverallontogeny, age states: primarymonocentrics(pl, j, im age states),polycentrics(v, gl-g3, ss age to (ss, s age states).In naturalhabitatsit is verydifficult states),secondarymonocentrics findsamplesof A. podagrariain earlystagesof development(pl-v),becausein shadethe species hardlyever reproducesfromseed. Plants flowerand bear fruitonly in sunlit placesin a forest,and evenhereyoungplantsdie in thesameyearthattheyemerge,being Seedlingsof withtheolderplantsforwaterand nutrients. unableto toleratecompetition A. podagrariaperishevenin felledareas,wherethisspeciesflowersand fruitsprofusely. The earlystages of its ontogenymay be foundundera forestcanopy onlyin well-lit places wherethe herb layer is disturbedby the activityof digginganimals. Further willbe determined bytheiractiveexpansion successoftheseyoungplantsofAegopodium plantsoccupyit. into open groundbeforeneighbouring Comparison of the ontogenyof Aegopodiumpodagrariawith that of other longrhizomedplants in which the structureof matureindividualsis clearlypolycentric (AppendixI) suggeststhatA. podagrariais a typicalmodelforthegroupas a whole. DISCUSSION and qualitative The age stateof an individualcan be definedbya complexof quantitative featuresare: nutritional whichare recognizablein thefield.The mostimportant features, juvenile or mature dependenceon the seed; the presenceor absence of embryonic, and structures;the abilityof an individualto reproduceor to propagatevegetatively; the relativeproportionsof livingand dead, active(growing)and passive(non-growing) plantparts.The main age statesof seed plantsmaybe characterizedas follows: have partial heterotrophic nutrition,fromthe subSeedlings(pl) characteristically stancesofthematernalplantstoredin theseed (especiallyin thecotyledons).Outwardly, structures: connectionto a seed.Thereare alwaysembryonic thisis seenin morphological cotyledons,primaryroot and primaryshoot. though Juvenile individuals(j) are structurally simple.Theyno longerhavecotyledons, found not have features Juvenile still be structures someembryonic plants present. may formof leavesand of shootand rootsystems. in matureplants,suchas different This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions 688 Age statesofplants Immatureindividuals(im) are characterizedby featurestransitional fromjuvenileto matureplants,in particularthe formof the leaves and root system.They may adopt anothergrowthformby developinglateralshoots,or by changingfroma monopodial to a sympodialconstruction. The primaryshoot,or partofit,however,is stillpresent, Virginileplants(v) beginto showthemainfeaturestypicalofmatureindividualsof de species,but reproductive organsare stillabsent.The distinctionl betweenimmatureand virginileplantsof some speciesmaybe difficult to make,sincebothage statescoverthe ofthepre-reproductive transition froma juvenileto a maturestructure. The differentiation or even impossible,because the structureof their period of some species is difficult leaves, shootsand root systemshardlychangesat thistime(e.g. Colchicumspeciosum -Shorina 1967). Some plant specieshave an immatureage-stateforonlya veryshort time. In some cases, therefore, an immatureage-stateis not distinguished (Zhukova 1961; Vorontzova1967,1968; Borissova& Popova 1971). of sexualorgansand bythe Reproductiveplantsare characterized bythedevelopment abilityto formseed. In youngreproductiveindividuals(gL), formationof new parts prevailsoverdeathof old parts,and is reflected in thebalance betweenlivingand dead structures. and fully-grown parts,and betweenactively-growing Matureindividuals(g2) showa relativeequilibriumin theprocessesof formationand death of structures. They usuallyshow the maximumyearlyincreasein biomass and maximumseed productivity. These are the individualswhichare at the peak of ontogeneticdevelopment.In old plants(g3), death of partsprevailsover the formationof newones,thereproductive is diminished, as is therateofrootand shootformation. activity is simplified Not infrequently plant structure (forinstancethe abilityto formrunners is lost,as in Mercurialisperennis). The reproductive period is subdividedmainlyaccordingto the balance betweenthe For severallifeformssome attempts processesof formationand death of structures. and have been made to get a quantitativeexpressionforthisbalance: forfirm-tussock loose-tussockgrasses,by comparingtheoveralldiameterof thetussockto the diameter of itslivingpart(Zhukova 1961; Borissova& Popova 1971; Kozhevnikova& Trulevich 1971; Egorova 1972); and forshort-rhizomed plantsby comparingthe algebraicmean of rhizomeincreasein lengthforyear'n' to themeanforyear'n + 1F(Smirnova1968). It is moredifficult to subdividethereproductive periodof long-rhizomed plants.Mature individualsof long-rhizomed plantsare systemsof partialtufts(or orthotropic shoots) rhizomes.Each of thesetuftsor shoots has its joined by plagiotropic(communicating) own lifenistory:it appears,develops,ages and dies off.The balance betweenyoungand old tufts(or shoots)determines the age stateof theindividualas a whole,and different valuesoftheindexa,. age statesoflong-rhizomed plantscan be characterized bydifferent The formationof each youngstructure (runneror youngtuft)rejuvenatesan individual is therejuvenation. as a whole: the morenew structures are formed,the moreeffective of the age indexa,, in individualsof This processis expressedby periodicalfluctuation thesameage statein long-rhizomed plants(Smirnova1976b;Lomakina 1972; Smirnova & Toropova 1974; Toropova 1975). ceases and fruitdevelopment In subsenile(ss) and senile(s) plants,flowering practically is simplified. Juvenileleaves appear once moreand growth and thevegetativestructure betweensubsenileand senileplantsare usuallyquantitaratedecreases.The differences tive. We know of no detailed classificationof ontogeneticdevelopmentin the English languageliterature. The phases of development-pioneer, building,matureand degene- This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions L. E. GATSUKet al. 689 rate(Watt 1947)-have been used to studyanotherlevel of plant systems,the patches or population;buttheage stateconceptis based on thedevelopment withina community however-for of an individualplant. It is true that these ideas are interconnected, plantsare dominantin the pioneerand buildingphases,old example,pre-reproductive phase. and senileplantsin thedegenerate in polycarpicplants,althoughin some cases All age statesare usuallydistinguishable period is not shown (some tree species); nor is therea postthe post-reproductive period some plants periodin monocarpicplants.Duringthe re'productive reproductive able to (Rabotnov 1950): a plantpotentially showintervalsbetweenflowering frequently flowermay not in fact formreproductiveshoots for several years. Some perennial secondarydormancy(Rabotnov 1950; Kurkin1976),thegrowth herbsmaydemonstrate absent. of visibleaerial organsbeingentirely phases may be takenas analogous to morphogenetic In annual plants,the different age states(Sharitz & McCormick1973). The developmentof clonal plantsmay by consideredfromtwo aspects: eitherfrom particuleand its eventualdeath,or fromseed to clone as a seed to vegetatively-derived whole(withall its particules)and its eventualdeath.In thelattercase, one mayspeak of 'complicatedontogeny'(Levin 1966)or about the 'greatlifecycle'(Smelov 1936). Hence as a whole(forexample,the one may distinguishage statesbothforclone development tussockof Deschampsiacespitosa)as well as forthe developmentof separateparticules. is frequentor intense,clone structure maybecomeso complicatedthat If rejuvenescence it is impossibleto speak about theage stateof thecloneas a whole. The age stateclassificationmay also be applied with some modificationto species propagation,suchas deciduousaxillarybuds whichhave specializedorgansofvegetative (Dentaria bulbiferaL.5 Ficaria vernaHuds.), filialbulbs (Gagea lutea (L.) Ker-Gawl., L.). This G. minima(L.) Ker-Gawl.),or adventitiousbuds on theroots(Ajugagenevensis has also been used to describethe ontogenyof Viola mirabilisL., which classification has root suckers(Smirnova& Kagarlitzkaya1972). To describeage statesin ontogeneticstudies,some additionaibiometricindications are also used, whichgive quantitativeestimatesto some morphological,anatomical, ecological and physiologicalfeatures-forexample,the size of an individualand its orders,general parts,the lengthof annual shoots,the numbersof roots of different biomass of an individual,the area of its leaf surface,the size of the leaf blades, the densityof stomata,the diameterof its shoot apex, etc. Such biometriccharacteristics of growthprocessesin ontomake it possibleto understandthe changein theintensity to estimatethe intensityof some physiologicalprocessessuch as geny,and indirectly and respiration.Direct quantitativedata for certainbiochemicalprophotosynthesis cessesin ontogenyare available,forexample,thecontentof RNA and albumennitrogen age states(Bylova & Grosheva Rupr. of different in individualsof Libanotisintermedia of individualsin a givenage statechange temporally 1973). Biometriccharacteristics ecologicalconditionsand (yearlyvariability)in a singlecoenosis,as well as in different economicmanagementof the coenosis (Vorontzova1968; Zaugolnova underdifferent 1968; Zhukova 1973). change uninterruptedly Unlike the qualitativeindices, quantitativecharacteristics duringontogeny.Changesin the majorityof themcan be representedby a unimodal curve (Fig. 5), maximumvalues being reachedin the maturereproductivestate (g2). (1975),thepeak occursmidwaybetweentheprevailing Accordingto Uranov'shypothesis an organismin thefirsthalfofitsontogenyand the of and assimilation by energy uptake This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions 690 Age statesofplants 0~~~~~~ 2 3 ~~~~~~~~1 4 5 6++ 0 E'20 1 2 345 6I E~~~~~~~~ 234 56 240 - FIG. 5. The change of some biometriccharacteristicsduring the ontogeny of Anabasis aphylla. (a) heightof shrub above ground; (b) diameter of shrub; (c) lengthof 'skeletal branch'; (d) diameter of the basal part of the root; I = immatureplants, 2 = virginile plants, 3 = young reproductive plants, 4 = mature reproductive plants, 5 = old reproductiveplants, 6 = senile plants. Vertical line is standard error. however,the prevailingloss of energyin the second half. For some characteristics, maximum maybe displacedto age statev,gl or g3 (Zaugolnova1968;Smirnova 1974). Field observationsand studieson biometriccharacteristics ofdifferent individualsof a their that state show some varies: individualsare given age developmentalcapacity others are individuals survive greatlysuppressed;large stronglydeveloped, throughall ones die without but a The state. age states, suppressed passing particularage prospects of developmentforthetwo typesof individualare different (Werner1975),so thatit is vitalityof individuals(Smirnovaet al. 1976). possibleto speak of thedifferent threemaintypesof ontogenyin polycarpicplants.The first It is possibleto distinguish of theindividualin ontogeny,and by lack of visibledisintegration typeis characterized of individualstructure lack of simplification at theend of ontogeny;thistypeis peculiar to themajorityof treesand to some speciesof tap-rootedsemi-shrubs (e.g. nos 1-11, 17, 27 and 28 in AppendixI). The second typeis characterizedby incompleteor partial oftheindividualplant,occurringusuallyat theend of ontogeny;thistype disintegration of developmentis typicalof tap-rootedsemi-shrubs and some perennialherbs(e.g. nos Plants 26 and 29-33 in 25, AppendixI). developingin thesetwo ways are vegetatively and of the renewal immovable, populationis possibleonlyby seeds. The thirdtypeof of the individualinto particules., is characterized by completedisintegration ontogeny in the first half of and whichmayoccurboth ontogeny at itsend; in thesenileage state, forexample).structural branching, maybe seen(no morethansecond-order simplification Such completedisintegration guaranteesvegetativepropagation,but it is not always sufficient to provideprolongedmaintenanceof the populationin the absence of seed Maintenanceofthepopulationbyvegetative reproduction. propagulesis possiblein two This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions 691 L. E. GATSUK etal. 80^ ct0 0 Q 0 o c " o 4 ;ON Q > > Eor r *^; = Y *~~~~~~C > o > b >-b >~~~~~~1- >b > 0,oo Q t. rc Cd . s< Cd~~S This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions 692 Age statesofplants cases: (a) if 'filial'individualsare rejuvenatedin comparisonwith'maternal'ones; or (b) if rejuvenescence does not occur,but insteadthe individualformsa clone in age statesi-g2. Thisthirdtypeof ontogenyhas threevariants(Table 2). Variety(3)a (Fig. 4(a), g2) has shootsin thematureage state,and is seenin tussockgrasses,sedges,and densely-packed in plantswithshortrhizomes,tuber-bulbs, and bulbs(e.g. nos 37-40,41-57, 76-80 and 81-88 in AppendixI). These plantsmay show vegetativepropagation.Varieties3(b) tuftsor shootsfrom and 3(b') are characterized by significant spacingof newly-formed one another;completedisintegration mayoccurbothearlyand late.Specieswiththistype of developmentare capable of spreadingwidelyand of propagatingvegetatively (longrhizomedand root-sucker plants,suchas nos. 58-71and 74-75in AppendixI). Rejuvenesbetweenvarieties(3)b and 3(b') is cence of 'filial'plantsis also possible.The difference that3(b) formspartialtuftsand 3(b') onlypartialshoots(Fig. 4(b)). These variedtypesof ontogenyare relatedto plantgrowthform,and in largedegree definetheage-statespectrum*of populationsand theplace of a speciesin thecoenosis. Numerouscoenopopulationsof different speciesin a rangeof ecological,geographical thesestudiesshowthesimilarities and successionalsituationshavenowbeeninvestigated; and differences of age-statespectraof particularplant growthforms(Zaugolnova & Smirnova1978).Variationsin age-statespectrain timeand space,and theconnectionof thesechangesto environmental changes,are known(Uranov et al. 1977). These studies giveus an insightintothemainfeaturesofpopulationstrategy of different plantspecies, featureswhichare important bothin theoryand in practice. ACKNOWLEDGMENTS The researchesdescribedhere were conductedat the Lenin State Teachers' Training Instituteof Moscow. Theywereinitiatedby thelate ProfessorA. A. Uranov,and from 1964 to 1974 were underhis constantconstructive leadership.The authorsthankall personswho helpedin fieldand laboratorystudiesand in preparingthispaper, V. N. Schukina,and especiallyDr J. WhitefortheEnglishtranslation. REFERENCES in steppegrasses.BotaniBorissova,I. 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Livingconditionsof species in plant associations.BulletenMoscovskogo obtshestva otdelbiologicheskij, prirody, 64, 77-92 (in Russian). ispytatelej This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions 694 Age statesof Dlants ofthephytocoenopopulation Uranov,A. A. (1975).Age spectrum as a function oftimeand energetic wave processes.Biologicheskie Nauki,2, 7-34 (in Russian). L. B., Smirnova, 0. V. (1977). Plant Coenopopulations (Development and Uranov,A. A., Zaugolnova, Interrelations). Nauka, Moscow(in Russian). Wardle,P. (1961). FraxinusexcelsiorL. BiologicalFlora of theBritishIsles.JournalofEcology,49, 739-751. Watt,A. S. (1947).Patternand processin theplantcommunity. Journal ofEcology,35, 1-22. P. A. (1975).Predictions Werner, offatefromrosettesize in teasel(Dipsacusfullonum L.). Oecologia, Berlin,20, 197-201. Zaugolnova,L. B. (1968). Age statesin the ontogenyof ash (FraxinusexcelsiorL.). Problemsof Morphogenesis ofFlowering PlantsandtheComposition oftheir Populations (Ed. byA. A. Uranov), pp. 81-102.Nauka,Moscow(in Russian). and dynamicsof coenopopulations of Zaugolnova,L. B. & Smirnova, 0. V. (1978). Age structure perennialplants.Zhurnalobscheibiologii,34, 849-858(in Russian). ofage changesintussockhairgrass. Zhukova,L. A. (1961).Thepeculiarities Morphogenesis ofPlants,2, 635-638(in Russian). of tuftedhairgrass(Deschampsiacaespitosa(L.) P.B.) Zhukova,L. A. (1973). Age-statecomposition oftheNorthDvina and theOka. Biologicheskie populationson water-meadows Nauki,7, 67-72 (in Russian). 6 November 1979) (Received *APPENDIX I No. theontogeny Authorwhodescribed Species I. Trees 1. BetulapubescensEhrh. Ldb. 2. B. tortuosa Ehrh. 3. B. verrucosa 4. FraxinusexcelsiorL. Bge. persicum 5. Haloxylon (Minkw.)Jljin 6. H. aphyllum Mill. 7. Malussylvestris 8. Picea excelsaL. Fisch.etMay 9. P. schrenkiana L. 10. Pinussylvestris 11. QuercusroburL. 1962 Serebrjakov 1962 Serebrjakov 1962 Serebrjakov Zaugolnova1968 1962 Serebrjakov 1962 Serebrjakov 1962 Serebrjakov 1962 Serebrjakov 1962 Serebrjakov 1962;Kravchenko1971 Serebrjakov 1962 Serebrjakov II. Shrubs avellanaL. 12. Corylus verrucosa Scop. 13. Euonymus 14. PrunusspinosaL. 15. Rosa albertiRgl. L. 16. R. spinosissima Domanskaja& Rodman1954 Serebrjakov, 1962 Serebrjakov 1962 Serebrjakov 1962 Serebrjakov 1962;Rodman1957 Serebrjakov III. Semi-shrubs Gatsuk& Vorontzova1976 17. AnabasisaphyllaL. & Chernyshova 1955 Serebrjakov 18. Andromeda polifoliaL. 1962 Zlobin1963;Serebrjakov (L.) Hull 19. Callunavulgaris Antonova1971 L. s.l. nigrum 20. Empetrum Gatsuk1968 Pall. 21. Hedysarumfruticosum & Chernyshova 1955 L. Serebrjakov myrtillus 22. Vaccinium & Chernyshova 1955 Serebrjakov 23. V.vitisidaeaL. * This listis incomplete, by collaboand includesonlythesespecieswhichhave beeninvestigated published. and accountsofthefindings ratorsoftheLeninMoscowStateTeachers'TrainingInstitute, Detailedreferences maybe obtainedon request. This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions L. E. GATSUK et al. No. Species theontogeny Authorwhodescribed IV. Low semi-shrubs Zaugolnova1974 Bge. diapensioides 24. Acantholimon Mikhailova1976 L. campestris 25. Artemisia Kozhevnikova& Trulevich1971 26. A. compactaFisch. Dorokhina1969 27. A. silvaticaMax. Zaugolnova1976 lenenseAdam. 28. Alyssum V. Herbs (a) Tap-rootedplants Bylova1974 L. 29. Buniasorientalis 1972 Piskovatzkova L. 30. Campanulaglomerata Bylova1968 31. Centaurea scabiosaL. 1967 Gulenkova1974;Mikhailovskaja L. 32. Lotuscorniculatus 1972 Blagoveschenskaja Knorr. 33. Lagochilusschugnanicus Bylova& Grosheva1973 Rupr. 34. Libanotisintermedia Pokrovskaja1974 L. 35. Trifoliumpratense Mussina1976 (L.) Scop. maculatus 36. Achyrophorus plants (b) Short-rhizomed Gulenkova1974 (Kramer)Garcke 37. Lathyruspannonicus Gulenkova1974 L. 38. L. sylvestris Gulenkova1974 L. 39. L. vernus Bakhmatova1975 Bernh. lobelianum 40. Veratrum (c) 'Bunchy'plants Kurchenko1975 L. 41. Agrostis stolonifera Kurchenko1972,1973 P. Smirn. 42. A. syreistschikowii Kurchenko1974 43. A. tenuisSibth. Kurchenko1967 Pall. ex Kunth 44. Alopecurus vaginatus Graschenkova 45. Carexcaryophyllea 1972 Latour. Fomichoeva1978 46. C. digitataL. Denissov1968 47. C. humilis Leyss. L. Egorova1972 48. Dactylisglomerata Zhukova1961 49. Deschampsia caespitosaBeauv. Mikhailova1977 50. FestucabeckeriHack. Ermakova1972 Huds. 51. F. pratensis Cheburaeva1974 (Huds.) Pilg. 52. Helictotrichon pubescens Cheburaeva1973 (Hack.) Kitagava 53. H. schellianum Bardonova1975 54. Hordeumbrevisubulatum (Trin.)Link 1941 Poshkurlat 55. Lasiagrostis (Trin.)Kunth. splendens Matveev1972 L. 56. Phleumpratense Bedanokova& Mikhailova1975 57. StipapennataL. Vorontzova, plants (d) Long-rhizomed Smirnova1967 L. 58. Aegopodiumpodagraria Smirnova1967 L. ranunculoides 59. Anemone L. Smirnova1967 60. Asarumeuropaeum Egorova1976 61. Bromusinermis Leyss. Smirnova1967 62. CarexpilosaScop. Lomakina1972 J.Gay 63. C. pachystylis Graschenkova 1974 64. C. praecoxSchreb. Willd. Zaugolnova1974 65. Galiumruthenicum L. Snagovskaja1965;Grigorjeva1974 66. Medicagofalcata Smirnova& Toropova1974 L. 67. Mercurialisperennis Donskova1968 68. 7rifolium Bieb. ambiguum & Kagarlitzkaja1972 Serebrjakova L. 69. Veronica longifolia & Kagarlitzkaja1972 Serebrjakova 70. V.incanaL. & Kagarlitzkaja1972 Serebrjakova 71. V.spicataL. 72. AsperulaodorataL. luteum Huds. 73. Galeobdolon plants (e) Stoloniferous Smirnova1974 Smirnova& Toropova1975 This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions 695 Age statesofplants 696 plants (f) Root-suckering Bylova1975 (L.) D.C. tataricum 74. Mulgedium Smirnova& Kagarlitzkaja1972 L. 75. Violamirabilis (g) Tuberousplants Beldeleva1972 (Steph.)Pers. bracteata 76. Corydalis Smirnova1974 etKoerte 77. C. cava(L.) Schweigg. Smirnova1974 78. C. halleriWilld. Smirnova1974 (L.) Merat 79. C. intermedia Smirnova1974 (Pall.) Pers. 80. C. marschalliana (h) Bulbousplants Shorina& Smirnova1976 L. 81. Alliumvictorialis Shorina& Smirnova1976 82. Gagealutea(L.) Ker-Gawl. Shorina& Smirnova1976 (L.) Ker-Gawl. 83. G. minima 1971 Shorina& Prosvirnina woronowii Losinsk. 84. Galanthus Smirnova1967 85. ScillasibiricaAndrevs. (i) Tuber-bulbousplants Shorina1968 Stev. 86. Colchicum speciosum Shorina1974 87. Crocusscharojanii Rupr. 88. C. vallicola Herb. Shorina 1974 This content downloaded from 134.129.182.74 on Thu, 19 Sep 2013 20:12:54 PM All use subject to JSTOR Terms and Conditions