全 文 ：中国藓类植物无性繁殖体的初步观察*
裴林英1,2,刘暋倩1,贾暋渝2**,王幼芳1
**
(1华东师范大学生命科学学院,上海暋200062;2中国科学院植物研究所
系统与进化植物学国家重点实验室,北京暋100093)
摘要:无性生殖在苔藓植物的生活史中起着重要的作用,并且常通过各种无性繁殖体来完成。无性繁殖体
的形态常被用来辅助鉴定一些不育的藓类植物。本文通过对38种藓类植物的无性繁殖体进行显微观察,
结果显示:无性繁殖体在不同的藓类植物之间已经过多次演化;无性繁殖体的形态在种内是相对稳定的,
且与其着生位置、配子体的分枝方式密切相关,而与植物的系统位置以及生境并无直接的关系;无性繁殖
体的颜色与其胞壁的厚度以及表面纹饰密切相关。此外,无性繁殖体的产生常常与假根、原丝体有共存关
系。在研究中发现,藓类植物的无性繁殖体主要包括原丝体芽胞、无性芽胞 (叶生芽胞、中肋芽胞、枝生
芽胞)、芽体、假根芽胞和假根状块茎;其中原丝体芽胞和无性芽胞最为常见。
关键词:藓类植物;无性繁殖体;原丝体芽胞;芽胞;芽体;假根芽胞;假根状块茎
中图分类号:Q944暋暋 暋暋 暋暋文献标识码:A暋暋暋暋暋暋暋文章编号:0253灢2700(2010)02灢103灢11
ObservationsontheAsexualDiasporesofMossesinChina
PEILin灢Ying1,2,LIUQian1,JIAYu2**,WANGYou灢Fang1**
(1SchoolofLifeScience,EastChinaNormalUniversity,Shanghai200062,China;
2StateKeyLaboratoryofSystematicandEvolutionaryBotany,InstituteofBotany,
ChineseAcademyofSciences,Beijing100093,China)
Abstract:Asexualreproductionplaysanimportantroleinthelifecycleofbryophytes.Usualy,mossescom灢
pletetheirasexualreproductionbydifferentdiaspores.Themorphologyofasexualdiasporehasbeencom灢
monlyusedfordelimitingsterilemosses.Inthepresentpaper,asexualdiasporesof38gemmiferousmosses
wereobservedunderlightmicroscope.Theresultsshowedthattheasexualdiasporeshaveevolvedseveral
timesindependentlyindifferentmosslineages.Themorphologyofasexualdiasporeisgeneralystablewithin
species,andiscloselyrelatedtoitsgrowingpositionandbranchingmodeofgametophyte,butnottophylo灢
geneticpositionsoftheplantsandtheirhabitats.Thecolorofdiasporeisdeterminedbythethicknessofthe
diasporalwalanditsornamentationonthesurface.Inaddition,theasexualdiasporesoftencoexistwithrhi灢
zoidsorprotonemata.Inthisstudy,asexualdiasporesincludeprotonemalgemmaeandgemmae(laminar
gemmae,costalgemmae,stemgemmae),bulbils,rhizoidaltubersandrhizoidalgemmae.Amongthem,
protonemalgemmaeandgemmaearethecommonestasexualdiaspores.
Keywords:Asexualdiaspores;Bulbils;Gemmae;Mosses;Protonemalgemmae;Rhizoidalgemmae;Rhi灢
zoidaltubers
暋暋Asexualreproductionplaysanimportant
roleinthelifecycleofbryophytes.Itispredom灢
inantinthebryophytesespecialywhenplants
areunderextremely hostileconditions (Jia,
云 南 植 物 研 究暋2010,32(2):103~113
ActaBotanicaYunnanica暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋DOI:10灡3724/SP灡J灡1143灡2010灡09221
*
**
Foundationitems:TheNationalNaturalScienceFoundationsofChina(Nos.30810103901and30770160)
Authorsforcorrespondence;E灢mail:yjia@ibcas灡ac灡cn;yfwang@bio灡ecnu灡edu灡cn
Receiveddate:2009灢11灢11,Accepteddate:2010灢02灢15
作者简介:裴林英 (1979-)女,博士研究生,主要从事苔藓植物学的分类学研究。
1994).Inmosses,asexualreproductioniscom灢
pletedbyvariousdiaspores,includingcaducous
shootapices,caducousbranchlets,flageliform
shoots,bulbils,caducousleaves,caducousleaf
apex,fragileleaves,rhizoidaltubersandgem灢
mae(ImuraandIwatsuki,1990;Newtonand
Mishler,1994;Imura,1994;Laaka灢Lindberg
etal灡,2003).Insomegenera,e灡g.,Bryum,
Campylopus,and Pohlia,asexualdiaspores
wereusedasoneofthediagnosticcharactersfor
specificidentification(CrundwelandNyholm,
1964;Syed,1973;Smith,1978;Mohamed,
1979;Shaw,1981;SlooverandSzmajda,1981;
Flowers,1983;Cetin,1999;AshtonandRaju,
2001;SherockandShaw,2005).
Duetoitsgreatvariabilityoftopography
andclimate,Chinaboastsarichmossfloraof
havingca.16% mossspeciesoftheworld(Red灢
fearnetal灡,1996).Thus,theknowledgeofdi灢
asporesofChinesemossfloraisimportantto
betterunderstandthesystematicrelationshipsa灢
mongthebryophytegroups.Theasexualdia灢
sporesofmossesinChina,however,havenot
beencomprehensivelyinvestigated,andtheter灢
minologyfordescribingthemisnotstandardized
andsometimesconfusing.Forexample,thea灢
sexualdiasporesofChinese mosses werenot
specifiedandwerecaledasgemmaeexceptfor
theflageliformbranchesinsomeChineselitera灢
tures(Gao,1994,1996;HuandWang,2005;
Li,2000,2006;WuandJia,2004;Wu,2002).
Anotherexampleisthatthebulbils,commonly
referringtothediasporeofBrachymeniumexile
bysomeauthors (Smith,1978;Lontonand
Schuster,1983;Reese,1984)werealsocaled
asgemmaebyLi(2006).
Inthepresentstudy,38gemmiferousmos灢
sesrepresenting12familiesand27generafrom
Chinawereexaminedunderthelightmicroscope
(LM).Ouraimsare1)todescribethegross
morphologyofasexualdiasporesfoundinsome
Chinesemosses,andtoclassifytheasexualdia灢
sporesintotypesaccordingtotheirgrowingpo灢
sitionanddevelopment;2)torecognizevarious
factorsthatafectthecharactersofasexualdiaspores
inmosses;3)todeterminethesignificanceofdi灢
asporalcharactersintaxonomyandsystematic
evolution.Thetermsconcerningasexualdia灢
sporesfolowImuraandIwatsuki(1990)and
NewtonandMishler(1994).
Materialsandmethods
Thirty灢eightgemmiferousspecimenswerethoroughly
cleanedthroughaseriesofrinsesandmountedinwater
(DuckettandPressel,2003)andphotographedwitha
digitalcameraunderOLYMPUSCX31lightmicroscopes
(LM).Vouchers(Table1)aredepositedinthefolowing
herbaria:herbariumoftheInstituteofBotany,Chinese
AcademyofSciences,Beijing(PE),herbariumofKun灢
mingInstituteofBotany,ChineseAcademyofSciences
(KUN),andherbariumofEastChinaNormalUniversity
(HSNU).
Classificationofasexualdiaspores
Charactersoftheasexualdiasporesinthe38species
ofgemmiferousmossesobservedinthepresentstudyare
summarizedinTable1.
Accordingtotheoriginoftheasexualdiaspores,
NewtonandMishler(1994)recognizedthattheasexual
diasporesmaycomeprimarilyfromtheasexualspores,
protonema,rhizoids,shoots,leaves,modifiedprotonema
andgametophyte.Inthisstudy,asexualdiasporesinclude
protonemalgemmae(fromprotonema),rhizoidaltuber
andrhizoidalgemmae (from rhizoids),bulbils (from
shoots),andgemmae(from modifiedprotonema).A灢
mongthem,protonemalgemmaeandgemmaearethe
commonesttypesofdiaspores.
1.Protonemalgemmae
Protonemalgemmaeusualyformedontheprotone灢
matawhichoccurinleafaxil.Theirshapesarediverse,
includingfilamentous(Tortulaschmidi,Plate栺:6),
clavate (Leptodontium flexifolium,Plate 栺:8)and
spherical(Barbulacoreensis,Plate栺:11-12).
2.Modifiedprotonema灢gemmaes.s.(probablyrepresen灢
tingaregeneratingprotonemainastateofarresteddevel灢
opment)
Gemmaearespecializedorgansservingasasexualre灢
production.Thegemmaeobservedwereclassifiedintothe
folowingthreecategoriesbasedondevelopmentalorigin:
laminargemmae,costalgemmaeandstemgemmae.
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暋暋1)Laminargemmae
Laminargemmaeareusualyclavateorfusiform,o灢
riginatedfromleafcels,e灡g.,inPlagiotheciumlatebri灢
cola(Plate栿:34)andHookeriaacutifolia(Plate栿:31
-32).
2)Costalgemmae
Costalgemmaeareusualyclavateorfilamentous,
andderivedfromcostacels.Theycanbeproducedonthe
costaattheleafapex,middleorbase,onventralordorsal
side,e灡g.,inSyrrhopodonparasiticus(Plate栺:1-2)
andMitthyridiumfasciculatum (Plate栺:4).
3)Stemgemmae
Stemgemmaeoftenoccuronstemsorshortbran灢
ches,derivingfromcorticalcels.Theymaybeclavateor
filamentous,e灡g.,inClastobryopsisplanula(Plate栿:
36)andTrachylomaindicum (Plate栻:28).
3.Bulbils
Bulbilsarehighlycondensed,vermicularorbud灢like
structures,withleafprimordialfoundinleafaxilofsome
mosses(NewtonandMishler,1994).Itisregardedasan
extremelyreducedbranch(ImuraandIwatsuki,1990).
Twokindsofshapesarefoundinthistypeofdiaspore:
bulk灢likebodies (e灡g.,in Brachymeniumexile,Plate
栻:20)andvermicularones(e灡g.,inPohliaproligera,
Plate栻:19).
4.Rhizoidaltubers
Rhizoidaltubersaredefinedasasexualorgansoccur灢
ringontherhizoids,oftensphericaltoelipsoidal,pyri灢
form,oroccasionalytuberous,thick灢waled,reddish
browntodarkbrown(Whitehouse,1966),e灡g.,inBry灢
umbornholmense(Plate栻:22).
5.Rhizoidalgemmae
Rhizoidalgemmaeareoftenfilamentous,branched,
orclavate,thick灢waledanddarkcolored,originateddi灢
rectlyfromrhizoids,e灡g.,inRhizomniumtuomikoski
(Plate栻:26).
Discussion
Themajoritymosseswithasexualdiaspores
inthisstudyaredioicousorsterileaspreviously
noted (Longton,1997;ImuraandIwatsuki,
1990).Thissuggeststhatasexualdiasporesare
foundmorefrequentlyindioicousorsterilemos灢
sestocompensateforsexualreproduction(Imu灢
ra,1994).
Thecharactersofasexualdiasporesaresta灢
blewithineachspecies,butnotdirectlyrelated
tothesystematicpositionsandhabitatsofthe
plants.Forexample,thegemmaeofHookeria
acutifoliaaresimilartothoseofPlagiothecium
latebricolainmorphology,butthetwospecies
arenotcloselyrelatedandwithdifferenthabi灢
tats.Forthisreason,itisclearthattheasexual
diasporeshaveevolvedseveraltimesindepend灢
entlyinthedifferentmosseslineages(White灢
house,1966).
Theoriginofdiasporeswasoftenusedfor
classifyingtheasexualdiaspores(Goebel,1905;
NewtonandMishler,1994).Inpresentstudy,
wefoundthatthecharactersofasexualdiaspores
areoftendependedontheirgrowingpositionson
theplants.Forexample,themorphologyofthe
costalgemmaearefairlyuniform,suchascla灢
vatetofilamentous,yelowishbrownandthin灢
waled.Inaddition,theasexualdiasporesfor
somespeciesareaffectedbyspecialenvironmen灢
talconditionsordifferentontogeneticphases.
Forexample,inBarbulahorrinervis(Plate栺:
13),theacanthoidstructureisproducedbythe
germinationofdiaspores.Moreover,thedia灢
sporesofPohliaflexuosaareoftenovateinthe
earlystageofdevelopment,butvermicularin
thelaterone(Li,2006).Onlytheovatedia灢
sporeswereobservedinourstudy (Plate 栻:
18).
Thebranchingmodesofgametophyteplay
animportantroleinasexualreproduction(Jia,
1994).Inthisstudy,theasexualdiasporesin
acrocarpusmossesareproducedonanypartsof
gametophyte,andtheir morphologicalcharac灢
tersarealsovaried.However,inpleurocarpous
mosses,theasexualdiaspores,onlyoccuringon
stems,protonemataorleaves,areoftenlinear
orclavate,yelowishbrown.
Thecolorsofasexualdiasporesaremainly
determinedbythecombinationofwalpigmen灢
tationandcopiousquantitiesofyelowtobrown
lipiddropletswithintheircels(ImuraandIwat灢
suki,1990;DuckettandPressel,2003).More灢
over,thecolorsofasexualdiasporesareoften
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dependedonthicknessofdiasporalwalsandits
ornamentationonthesurface.Thethick灢waled
diasporesareoftenreddishbrownoryelowish
brown,verruculoseorconvexonthesurface,
butthethin灢waledonesarelightyelow,hya灢
lineandsmoothonthesurface.
Rhizoidsorprotonemaareusualyproduced
onleaves,costaeorthebasesofplantsasthea灢
sexualdiaspores.Theprotonemaareoftenir灢
regularly branched or stalk灢like, yelowish
brown,andoccurontheleafaxil,e灡g.,inBar灢
bulaindica(Plate栻:16).Therhizoids,how灢
ever,oftenoccurontheleafapice,costaeorthe
basesofplants.Therhizoidsontheleafapex
andcostaaresimilartonormalonesinmorphol灢
ogy,e灡g.,in HookeriaacutifoliaandMitthy灢
ridiumfasciculatum .Whereas,therhizoidson
the base of plants are usualy irregularly
branched,verruculoseonthesurface,e灡g.,in
BryumbornholmenseandPohlialutescens.It
isnoteworthythattheasexualdiasporesonthe
upperpartofstemrarelycoexistwithrhizoidsor
protonema,butdirectlyclusteronstems,e灡g.,
inClastobryopsisplanula.
Mostly,mossescompleteasexualreproduc灢
tionbyasingletypeofdiaspore.Onlyafewof
mossesproducetwoormoretypesofdiaspores
(Whitehouse,1966).Inpresentstudy,two
typesofgemmaeoccurondifferentpositionsin
Bryumcapilare.Onetypeofgemmaeoccurs
ontheyelowishbrown,tuberculate,under灢
groundrhizoids.Thegemmaeareoftendecidu灢
ous,multicelular,pyriformtonearlyspherical,
reddishbrown,withthickandsmoothwals,
andtheyareusualyinflated,ca.100-120毺m
long,65-70毺mwide.Thesecondtypegrows
onthestemneartheleafaxil.Thegemmaeare
uniseriate(upto25cels),linear,yelowish
brown,thick灢waled,verruculoseonsurface,
andca.1灡1mmlong.
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ExplanationofPlate
Plate栺暋1-9.Costalgemmaeandprotonemalgemmae暋1,2.
CostalgemmaeofSyrrhopodonparasiticus;3.Costalgemmaeof
Calymperesafzeli;4.CostalgemmaeofMitthyridiumfascicula灢
tum,showingrhizoids(atarrow);5.Protonemalgemmaeof
Anoectangiumfauriei,showingprotonemata (atarrow);6.
ProtonemalgemmaeofTortulaschmidi;7.Protonemalgem灢
maeofDesmatodongemmascens,showingprotonemata(atar灢
row);8.ProtonemalgemmaeofLeptodontiumflexifolium;9.
ProtonemalgemmaeofDidymodonrigidulus,showingprotone灢
mata(atarrow).10-15.ProtonemalgemmaeandRhizoidaltu灢
bers暋10.ProtonemalgemmaeofDidymodonferrugineus;11,
12.ProtonemalgemmaeofBarbulacoreensis,showingprotone灢
mata(atarrow);13.ProtonemalgemmaeofBarbulahorrinervis,
showinggerminatedgemmae(atarrow);14.Protonemalgemmae
ofBarbulaamplexifolia;15.ProtonemalgemmaeofTimmiela
anomala
Plate栻暋16-18.ProtonemalgemmaeandRhizoidaltubers暋
16.ProtonemalgemmaeofBarbulaindica;17.Rhizoidaltubers
ofPohlialutescens,showingrhizoids(atarrow);18.Protone灢
malgemmaeofPohliaflexuosa.19-27.Protonemalgemmae,
bulbils,rhizoidaltubers,stemgemmaeandrhizoidalgemmae暋
19.BulbilsofPohliaproligera;20.BulbilsofBrachymenium
exile;21.BulbilsofBryumgemmigerum;22.Rhizoidaltubers
ofBryumbornholmense;23.RhizoidaltubersofBryumthom灢
soni,showingrhizoids(atarrow);24.Rhizoidaltubersand
stemgemmaeofBryumcapilare(Ashowsstemgemmae,B
showsrhizoidaltubers);25.StemgemmaeofBryumtortifolium;
26.RhizoidalgemmaeofRhizomniumtuomikoski;27.Proto灢
nemalgemmaeofRhachitheciumperpusilum,showingprotone灢
mata(atarrow).28-30.Protonemalgemmae,laminargemmae
andstemgemmae暋28.StemgemmaeofTrachylomaindicum;
29.ProtonemalgemmaeofPterobryopsisorientalis;30.Proto灢
nemalgemmaeofHorikawaeanitida
Plate栿暋31-36.Protonemalgemmae,laminargemmaeandstem
gemmae暋31,32.Laminargemmaeof Hookeriaacutifolia,
showingrhizoids (at32);33.ProtonemalgemmaeofCya灢
thophorelahookeriana;34.LaminargemmaeofPlagiothecium
latebricola,showingrhizoids(atarrow);35.Stemgemmaeof
Pylaisiadelphayokohamae;36.StemgemmaeofClastobryopsis
planula.37-42.Stemgemmaeandbulbils暋37.Stemgemmae
ofClastobryopsisrobusta;38.StemgemmaeofClastobryum
glabrescens;39.StemgemmaeofGammielatonkinensis;40.
Bulbilsof Pseudotaxiphylum pohliaecarpum;41,42.Stem
gemmaeofIsopterygiumpropaguliferum
011暋暋暋暋暋暋暋暋暋暋暋暋 暋暋暋暋暋暋暋云暋南暋植暋物暋研暋究暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋第32卷
暋裴林英等:图版栺暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋PEILin灢Yingetal灡:Plate栺
1112期暋暋暋暋暋暋PEILin灢Yingetal灡:ObservationsontheAsexualDiasporesofMossesinChina暋暋暋暋 暋暋暋
暋裴林英等:图版栻暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋PEILin灢Yingetal灡:Plate栻
211暋暋暋暋暋暋暋暋暋暋暋暋 暋暋暋暋暋暋暋云暋南暋植暋物暋研暋究暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋第32卷
暋裴林英等:图版栿暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋PEILin灢Yingetal灡:Plate栿
3112期暋暋暋暋暋暋PEILin灢Yingetal灡:ObservationsontheAsexualDiasporesofMossesinChina暋暋暋暋 暋暋暋