Download Age States of Plants of Various Growth Forms: A Review

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 .
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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
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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
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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.
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678
Age statesofplants
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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
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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
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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
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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
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L. E. GATSUK
et a!.
683
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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
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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
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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
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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
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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-
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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
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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
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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
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0,oo
Q t.
rc
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.
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Cd~~S
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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. V. & Popova,T. A. (1971). Age phasesof bunch-formation
cheskijzhurnalSSSR, 56, 619-626.(in Russian).
derPflanzen.Jena.
BoysenJensen,P. (1932). Die Stoffproduktion
characteristics
N. P. (1973). Morphologicaland physiological-biochemical
Bylova,A. M. & Grosheva,
zhurnal
SSSR, 58, 1510-1515(inRussian).
Rupr.Botanicheskij
ofage statesofLibanotisintermedia
L.) in thefloodmeadowsof
Egorova,V. N. (1972).The lifecycleoforchardgrass(Dactylisglomerata
77, 4,
prirody,otdelbiologicheskij,
the Oka river.BulletenMoscovscogoobschestva
ispytatelej
118-129(in Russian).
the importance
of the
in plantcommunities:
Grubb,P. J. (1977). Maintenanceof species-richness
niche.BiologicalReviews,
52, 107-145.
regeneration
Nauka,Moscow(in Russian).
inSilviculture.
Gupalo,P. I. (1969).AgeStatesandtheirImportance
ofplants.AnnualReviewofEcologyandSystemaHarper,J. L. & White,J. (1974).The demography
tics,5, 419-463.
and DynamicPlantEcology,2ndedn. Arnold,London.
K. A. (1973). Quantitative
Kershaw,
N. V. (1971). Dry Steppesof InnerTien-Shan.Ilim, Frunze(in
N. D. & Trulevich,
Kozhevnikova,
Russian).
* The relativeproportions
(Uranov1960).
ofeach age statein a coenopopulation
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.
693
Krenke,N. P. (1940). The Theoryof CyclicAgeingand Rejuvenation
ofPlantsand itsPracticalUse.
Selkhozgiz,Moscow (in Russian).
Kurkin,
K. A. (1976).SystemsResearchesofMeadowDynamics.Nauka, Moscow(in Russian).
Levin,G. G. (1966). Age changesin plants.Botanicheskij
zhurnalSSSR, 51, 1774-1795(in Russian).
Leysle,F. F. (1949). The ecologyand anatomyof halophytesand xerophytes
withreducedleaves.
zhurnalSSSR, 34, 253-266.(in Russian).
Botanicheskij
Lomakina,G. A. (1972). Some age statesof Carexpachystylis
Gay. Biologicheskie
nauki,3, 65-72(in
Russian).
Passecker,F. (1977). Theorie der ontogenetischen
Evolutionund AlterungholzigerGewachse.
Bodenkultur,
28,277-294.
Persikova,
Z. I. (1959).ThegreatlifecycleofDeschampsia
caespitosaL. Uchoenyje
zapiskiMoskovscogo
gosudarstvennogo
pedagogiclieskogo
instituta,
t. C, 5, 111-150(in Russian).
Rabotnov,T. A. (1950). The lifecycleof perennialherbaceousplantsin meadowcoenoses.Trudy
Botanicheskogo
instituta
AkademiiNauk SSSR, seria3, 6, 7-204 (in Russian).
determination
in herbage.Field Geobotany.
Rabotnov,T. A. (1960). Methodsof age and longevity
AkademiaNauk SSSR, Moscow (in Russian).
T. A. (1969). Some problemsof thestudyof coenoticpopulations.BulletenMoscovskogo
Rabotnov,
obschestva
ispytatelej
prirody,
otdelbiologicheskij,
74, 1, 141-149(in Russian).
Rabotnov,T. A. (1974). The Studyof Meadows.Moscovskijgosudarstvennyj
universitet,
Moscow
(inRussian).
Robbins,W. I. (1957). Physiologicalaspectsof ageingin plants.American
Journalof Botany,44,
289-294.
in woodyplants
de Muckadell,M. (1959).Investigation
Schaffalitzky
on ageingof apical meristems
and itsimportance
in silviculture.
i Danmark,4, 307-455.
ForstligeForsogsvacsen
Scharitz,R. R. & McCormick,J. F. (1973). Populationdynamicsof two competingannual plant
species.Ecology,54,723-739.
Schitt,P. G. (1958). Studyof the Growthand Development
of Fruitand BerryPlants.Selkhozgiz,
Moscow(in Russian).
ofthegrasslayerin oakwoods.Uchoenyje
Shik,M. M. (1953).Seasonaldevelopment
zapiskiMoscovskogogosudarstvennogo
pedagogicheskogo
instituta,
caphedrabotaniki,
73,2, 159-250(in Russian).
Shorina,N. 1. (1967).Lifecycleof Colchicum
Stev.in theforestand subalpinebeltsof west
speciosum
Transcaucasus.Ontogeny
andAge-StateComposition
ofFlowering
PlantPopulations
(Ed. byA. A.
Uranov),pp. 70-99. Nauka, Moscow (in Russian).
I. G. (1962).EcologicalMorphology
Serebrjakov,
ofPlants.Vysshajashkola,Moscow(in Russian).
E. W. (1960).PlantMorphogenesis.
Sinnott,
McGraw-Hill,New York.
Smelov,S. P. (1936).New trendsin thedevelopment
ofthetheoretical
ofthecultivation
of
principles
meadowsand pastures.Problemy
zhivotnovodstva,
11, 19-28(in Russian).
0. V. (1967). Ontogenyand age statesof Carexpilosa Scop. and Aegopodium
Smirnova,
podagraria
L. Ontogeny
PlantPopulations
andAge State Composition
ofFlowering
(Ed. by A. A. Uranov).
pp. 100-113.Nauka, Moscow(in Russian).
0. V. (1968).Some pecularities
ofvegetatively-movable
Smirnova,
plantlifecycles.Uchoenyje
zapiski
Permskogo
pedagogicheskogo
instituta,
64, 153-158(in Russian).
0. V. (1974).The characteristics
of vegetative
Smirnova,
reproduction
byherbaceousspeciesof oakwoodsin relationto populationmaintenance.
PlantPopulaAge-StateComposition
ofFlowering
tionsin connection
withOntogeny
(Ed. byA. A. Uranov),pp. 168-195.Moscow (in Russian).
Smirnova,0. V. & Kagarlitzkaja,T. N. (1972). On the two typesof lifecycleof Viola mirabilis.
zhurnalSSSR, 57, 481-492(in Russian).
Botanicheskij
0. V. & Toropova,N. A. (1974). The lifecyclesand age-statecompositionof somelongSmirnova,
in
PlantPopulations
rhizomedplantpopulationin oakwoods.Age-StateComposition
ofFlowering
connection
withOntogeny
(Ed. by A. A. Uranov),pp. 56-69.Moscow(in Russian).
0. V., Zaugolnova,L. B., Ermakova,
Smirnova,
I. M. (1976). Plant Coenopopulations
(Basic Concept
and Structure).
Nauka, Moscow(in Russian).
indicesin defining
Toropova,N. A. (1975). The use ofquantitative
age stagesin rhizomatous
grasses.
zhurnalSSSR, 60, 1297-1304(in Russian).
Botanicheskij
of a southernsemi-desert
L. I. (1967). Changingvitality
Vorontzova,
speciesas a resultof ecological
and Age-StateComposition
Plant Populations(Ed. byA. A.
conditions.Ontogeny
of Flowering
Uranov),pp. 132-154.Nauka, Moscow (in Russian).
and age-state
ofFestucasulcatapopulations
Vorontzova,
L. I. (1968).Changeofnumbers
composition
in different
conditionsof southern
semi-desert.
Uchoenyje
zapiskiPermskogopedagogicheskogo
instituta,
64,79-84(inRussian).
Uranov,A. A. (1960). 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.
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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
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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
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