全 文 ：广 西 植 物 Guihaia　Oct．２０１５,３５(５):６９７－７０３　　　　　　　　　　 http://journal．gxzw．gxib．cn　
DOI:１０．１１９３１/guihaia．gxzw２０１３１１０３９
申琳,张泽悠,夏乔莉,等．基于光合参数探讨四种藓类作水族箱植物的应用潜力[J]．广西植物,２０１５,３５(５):６９７－７０３
ShenL,ZhangZY,XiaQL,etal．PotentialoffourmossesasaquariumplantsＧdeducedfromtheirphotosyntheticparametersinwater[J]．Guihaia,２０１５,
３５(５):６９７－７０３
PotentialoffourmossesasaquariumplantsＧdeduced
fromtheirphotosyntheticparametersinwater
SHENLin,ZHANGZeＧYou,XIAQiaoＧLi,GUOShuiＧLiang∗
(CollegeofLifeandEnvironmentalSciences,ShanghaiNormalUniversity,Shanghai２００２３４,China)
Abstract:Aquaticmossescouldbeusedasaquariumplants,manysemiＧaquaticmossesarealsoabletogrowinaquarＧ
ia．IneasternChina,therearefewaquaticmosses．Isitpossibletousesometerrestrialmossesinaquaria?Inorderto
answerthequestion,weelucidatedtheadaptabilityoffocalterrestrialmossestowaterenvironment．Inthepresent
work,wemeasuredthephotosyntheticparametersoffourterrestrialmossesincludingBrachytheciumprocumbens,
Hypnumhamulosum,Leucobryumglaucum,andHedwigiaciliataunderconditionssimilartotheirnaturalhabitats
andthoseaftertheirsubmersioninwater．WealsomadetheirphotosyntheticlightＧresponsecurvesbyusingrectanguＧ
larhyperbolicmodel．Wefoundsignificantdiferencesamongtheirmaximumnetphotosynthesisrate(Pn),lightsatＧ
urationpoint(LSP),andlightcompensationpoint(LCP)．ThevariationrangesoftheirmaximalPn,LSPandLCP
werefrom１２２．５７５to１９．０９９μmolCO２􀅰kgＧ１DW􀅰sＧ１,from１１６６．００to６７０．０３０μmol􀅰mＧ２􀅰sＧ１,andfrom８５．０００
to５．３μmol􀅰mＧ２􀅰sＧ１,respectively．AfterBrachytheciumprocumbens,HedwigiahamulosumandLeucobryum
glaucumhadbeensubmergedinwaterfor３０d,theirmaximalPnwere１１０．７８％,８０．８４％and１０９．６３％oftheconＧ
trol,respectively,indicatingthatthesethreemossesareabletosurviveinwaterduringtheexperimentalperiod．While
submergedinwaterfor２０d,Hedwigiaciliatahadonly５．２５％netphotosyntheticrateofthecontrol,revealingthat
H．ciliateisnotabletogrowinaquaticenvironment．Wealsodiscussedtherelationshipsoftheirphotosynthesiswith
theirmorphologicalstructureandhabitatconditions．OuranalysesshowedthatBrachytheciumprocumbens,H．hamＧ
ulosumandLeucobryumglaucum,thoughdistributeinterrestrialhabitatsinthefield,werethreepotentialaquarium
plants．
Keywords:terrestrialmosses;photosynthesis;submersion;aquariumplants
CLCnumber:Q９４５．１１　　Documentcode:A　　ArticleID:１０００Ｇ３１４２(２０１５)０５Ｇ０６９７Ｇ０７
基于光合参数探讨四种藓类作水族箱植物的应用潜力
申　琳,张泽悠,夏乔莉,郭水良∗
(上海师范大学 生命与环境科学学院,上海２００２３４)
摘　要:水生藓类植物适宜作为水簇箱植物,许多半水生藓类植物同样能够生长于水体环境中.中国东部地区
的水生藓类植物种类不多,陆生藓类植物能否应用于水族箱中? 为了回答这一问题,需要阐明陆生藓类植物对
水体环境的适应能力.该研究测定了匐枝青藓(Brachytheciumprocumbens),弯叶灰藓(Hypnumhamulosum)、
白发藓(Leucobryumglaucum)和虎尾藓(Hedwigiaciliata)在与它们的自然生境相似条件下以及沉水环境下的
光合参数,并应用直角双曲线模型拟合了它们的光－光合响应曲线.结果表明:这四种藓类植物在最大净光合
收稿日期:２０１４Ｇ０６Ｇ２３　　修回日期:２０１４Ｇ０９Ｇ１８
基金项目:上海市科委项目(１２４９０５０２７００,１１３９１９０１２００);上海市教委大学生创新项目(BＧ７０６２Ｇ１２Ｇ００１０８１).
作者简介:申琳(１９９３Ｇ),黑龙江哈尔滨人,上海师范大学生命与环境科学学院２０１１级学生,(EＧmail)１０００３２９７９７＠smail．shou．edu．cn.
∗通讯作者:郭水良(１９６４Ｇ),教授,主要从事生态学教学科研工作,(EＧmail)gsg＠shnu．edu．cn.
速率(Pn)、光饱和点(LSP)和光补偿点(LCP)上存在很大差异.它们的最大净光合速率、光饱和点和光补偿点
的变异范围分别为１２２．５７５~１９．０９９μmolCO２􀅰kgＧ１DW􀅰sＧ１、１１６６．００~６７０．０３０μmol􀅰mＧ２􀅰sＧ１和８５．０００~５．３
μmol􀅰mＧ２􀅰sＧ１.在沉水环境中生长３０d后,匐枝青藓、弯叶灰藓和白发藓的最大净光合速率分别是对照的
１１０．７８％、８０．８４％和１０９．６３％,说明在实验周期里这三种藓类植物能够在水体环境中生存,而虎尾藓在水体中浸
泡２０d后,其最大净光合速率仅为对照的５．２５％,反映出该种植物并不适应水体环境.综上可知,四种藓类植物
的光合速率与其形态结构和原生境条件有很大的关系,虽然匐枝青藓、弯叶灰藓和白发藓主要分布于陆生环境,
但作为水族箱植物也具有一定的应用潜力.
关键词:陆生藓类;光合作用;沉水环境;水族箱植物
　　Aquaticmosseshavebeenappliedasdecorative
plantsinaquaria．Theyalsoprovideoxygen,hiding
places,andeggＧlayingsubstratesforfishes (Benl,
１９５８;Takakietal．,１９８２)．Inmarkets,therepresentaＧ
tiveaquariumbryophytesareFissidensfontanus,BryＧ
um pseudotriquetrum,Fontinalisantipyretica,LepＧ
todictyumriparium,PlatyhypnidiumriparioiＧdes,
Ricciafluitans,Ricciocarposnatans,TaxiphylＧlum
barbieri,andVesiculariadubyana (Benl,１９５８;Takaki
etal．,１９８２;Gradsteinetal．,２００３;Tanetal．,２００４)．
Mossesarefundamentalysimilartootherplants
intheirbasicnutritionrequirements．However,they
havespecificwaytoobtainnutrients．Evenifmosses
areabletousetheirrhizoidstogathersomenutrients,
theirrhizoidscannotpenetrateintosoils．Mosses
mainlyrelyonnutrientsfromdustontheirsurfacesor
dissolvedinrainfal,whichisaquitediferentstrategy
fromvascularplants．Typicaly,mosseshaveleavesof
onlyonecellayerinthicknessandwithoutcuticleand
specialprotectionstructures,exposingeveryleafcel
directlytotheirsurroundingstogetnutrients．ThegaＧ
metophytesofmanymossesareofshapeofsheets,filＧ
aments,twigs,etc．,withrelativelylargeleafsurfacearＧ
ea．Therefore,comparedwithmesicorxerictracheoＧ
phytes,mossesaremorphologicalysimilartoaquatic
tracheophytes．FromevolutionaryviewsandrecapituＧ
lationlaw,bryophytesbelongtoacladevolvedfrom
aquatictoterrestrialtaxa．Therefore,terrestrialor
semiＧaquaticmossesmaybeeasierintroducedintoand
adaptedtoaquaticenvironments．Forexample,Bryum
pseudotriquetrum,aspeciesgrowingonthinsoilsof
rocks,hasbeenfoundintheAntarcticdeepＧwaterarea
(Wangeretal．,２００６)．Fissidensfontanus,amossspeＧ
ciesoriginatingfromNorthAmerica,usualygrowson
rocks,treetrunks,andisalsoabletogrowinshalow
water(Crumetal．,１９８１),nowitisculturedasa
aquariumplantinSingapore(Tanetal．,２００４)．
IneasternChinaalongthePacificocean,thereis
fewaquaticbryophytes(Xu,１９８９;Liuetal．,２００５)．Is
itpossibletofindsometerrestrialbryophytesasaquarＧ
iumplants?Inordertoanswerthequestion,weconＧ
ductedexperimentstotesttheadaptabilityofsometerＧ
restrialbryophytespeciestowaterenvironment．
Brachythecium procumbens,Hypnum hamuloＧ
sum,LeucobryumglaucumandHedwigiaciliataare
fourabundantandwidelydistributedterrestrialmoss
speciesineasternChina．Insomestands,BrachytheciＧ
um procumbens and Hedwiagiahamulosum often
coverwideareas,appearas“greencarpet”,andH．cilＧ
iataappearsas“greypatch”onstonesandboulders,
whileLeucobryumglaucumas“whitegreenishpatch”
onforestfloorortrunkbaseofPinusmassonianaAsＧ
sociation．ThesefourmossspeciesarevaluableasorＧ
namentalplants．However,theiradaptabilityinaquatic
environmentisnotclear．
Photosyntheticparametersareimportantreference
valuesindicatingtheirabilitytoadapttoenvironments．
Therehavebeenconsiderablereportsaboutbryophyte
photosynthesis(Liuetal．,２００１;VanGaalenetal．,
２００７;Gofinetetal．,２００８)．Wateravailabilityisoneof
themostimportantfactorsthatlimitdistributionand
productivityofbryophytes．Dilksetal．(１９７５)found
thattherewasaspecificrangeofwatercontentforxeＧ
ric mossestokeepnormalphotosyntheticactivity．
Generalyspeaking,mossphotosynthesisrateisposiＧ
tivelycorrelatedwithmoisturecontentofambientenＧ
vironment．However,ifaboveorbelowacertainrange
ofwatercontent,thenetphotosyntheticrateofmost
mosseswouldbeinhibited．Liuetal．(２００１)found
thattheoptimummoisturecontentofThuidiumcymＧ
８９６ 广　西　植　物　　　 　　　　　　　　　　　　　　３５卷
bifoliumandChrysocladiumretrorsum was２００％to
４００％oftheirdryweightfortheirphotosynthesis．For
thebryophytes mainlygrowinginhumidenvironＧ
ments,themoisturecontenthasnosignificantlyefects
on net photosyntheticrate,such as Hylocomium
splendensinB．S．G．andPleuroziumschreberi(Busby
etal．,１９７８)．Thoughtherearemanyreportsaboutthe
toleranceofmossestodroughtbasedontheirphysioＧ
logicalresponsesincludingphotosyntheticandchloroＧ
phylfluorescenceparameters(Kalaposetal．,２００１,
Zhangetal．,２０１１),littleworkaboutthetoleranceof
terrestrialmossestosubmergedenvironmenthasbeen
conducted．
ThepurposeofthisworkistocomparetheadaptＧ
abilityoftheabovefourmossspeciestowatervia
measuringtheirphotosyntheticcapacityinwater,toeＧ
lucidatethepossibilitytoapplywaterＧtolerantterresＧ
trialmossesasaquariumplants．
１　MaterialsandMethods
Experimentalsamplesoffourmossspecieswere
colectedfromeasternChinaonMarch１２－１３,２００９．
Amongthem,Leucobryum glaucum wascolected
fromNanmingMountaininthesuburbofLishuicity
inZhejiang(２８°２６′０５″N,１１９°５４′１０″E,elevationca．８２
m),Hedwigiahamulosumfromnorthernsuburbof
Jinhuacity,Zhejiang(２９°１３′１９″N,１１９°３７′５６″E,elevaＧ
tionca．１１８０m),H．ciliatafromthesuburbofJinＧ
huacity,Zhejiang(２９°１２′５５″N,１１９°３８′２６″E,elevation
ca．１０９０m),andBrachytheciumprocumbensfromthe
BotanicalGarden,XuhuiCampusofShanghaiNormal
University(３１°０９′５１″N,１２１°２４′５０″E,elevation３m)．
Plantmaterialswereconfirmedunderamicroscope,
sporophytesandimpuritieswereremovedfromthe
samples．Thematerials (greengametophytes)were
washedthreetimeswithdistiled water,then were
driedbynaturalventilationforexperiment．
Aweekafterthecolection,３gdrymaterialwere
putintoanylonbag(１２cm×６cm,withmeshof５
mmindiameter)tomakeamossbag,atotaloffive
mossbagsweremadeforeachmossspecies,themoss
bagswereimmersedinaplasticbucket(height１８cm
×diameter２０cm)withtapwater(１０cmindepth)．
Duringtheexperiment,thesampleswerekeptatthe
temperatureof１２－２２℃andnotpumpedoxygeninto
thewater．Afterbeingsubmergedinwaterfor５,１０,
１５,２０,３０d,themossbagswereremovedfromwater,
respectively．Fivesamples(eachwithca．０．２gfresh
weight)weretookforeachspeciesasfiveduplicates．
Thematerial(ascontrol)wassoakedintotapwater
foroneminutebeforethedeterminationofitsphotoＧ
synthesisrate．Afterthemeasurement,thematerials
weredriedat８０℃for８handthenweighed．
Thenetphotosyntheticrate(Pn)offourmoss
specieswasmeasuredwithaportablephotosynthesis
system(GFSＧ３０００,Walzcompany,Germany)from９:
３０amto１１:００am,withtheleavecuvettecondition
beingsetwithrelativehumidityof(６０±１０)％,temＧ
peratureof２０℃andCO２concentrationof３４０μmol􀅰
molＧ１．PARwasgivenbyLEDLightSource３０４０ＧLof
theGFSＧ３０００atdiferentintensities．DatawererecorＧ
dedthreetimesforeveryPARintensity．
ThephotosyntheticlightＧresponsecurves were
modeledusingrectangularhyperbolicmodelasPn＝
a􀅰b􀅰PAR/(a􀅰PAR＋b)Ｇc,herePnisnetphotoＧ
syntheticrate,PARisphotosyntheticactiveradiation
measuredintheupperpartofthecuvetteofthestandＧ
ardmeasuringhead３０１０ＧS,a,bandcareparameters．
BasedonphotosynthesisＧlightresponsecurveequation,
LSPcorrespondingto９５％ofmeasuredmaximalPn
andLCPwerecalculated．
Althetreatmentswerereplicatedfivetimes．The
datapresentedarethemeans±SE．OneＧwayANOVA
wasemployedtotestthediferencesofthedatafrom
theexperimentswiththeprocedureofSPSS１１．０staＧ
tisticalpackage(SPSSCorp)．
２　ResultsandAnalysis
ThephotosyntheticlightＧresponsecurves,maxiＧ
malPn,LSPandLCPoffourmossspeciesundernatＧ
uralconditionarelistedinTable１．
AccordingtotheirmaximalPnfromhightolow,
fourmossspeciesarerankedasB．procumbens,HedwiＧ
giahamulosum,Leucobryumglaucum andHedwigia
９９６５期　　　　　　　申琳等:基于光合参数探讨四种藓类作水族箱植物的应用潜力
ciliata,theirmaximalPn (μmolCO２􀅰kgＧ１ DW􀅰sＧ１)
being１２２．５８,８７．２４,４１．１０and１９．１０,respectively;HedＧ
wigiaciliata hasthehighestLSP,being１１６６．００
μmol􀅰mＧ２􀅰sＧ１,thenB．procumbensandH．hamuloＧ
sum,being１０１８．６０and９４１．８０μmol􀅰mＧ２􀅰sＧ１,respecＧ
tively,andLeucobryumglaucumhasthelowestLSPat
６７０．００μmol􀅰mＧ２􀅰sＧ１;AsforLCP (μmol􀅰mＧ２􀅰sＧ１)
fromhightolow,fourspeciesarerankedasHedwigia
ciliata (８５．００),Brachytheciumprocumbens (５６．８０),
Hedwigiahamulosum (２０．５０)andLeucobryumglauＧ
cum (５．３)．
PnofBrachytheciumprocumbensafterimmerＧ
sioninwater:Afterbeingsubmergedinwaterfor５d,
thePnofB．procumbensat１６００μmol􀅰mＧ２􀅰sＧ１is
１２７．１０μmolCO２􀅰kgＧ１DW􀅰sＧ１,being９８．７４％ofthe
control,whichisnotsignificantlydiferentfromthe
control．Afterbeingsubmergedinwaterfor１０,１５and
２０d,itsnetphotosyntheticratesat１６００μmol􀅰mＧ２􀅰
sＧ１decreasetosomeextent,being７８．２１％,８７．７７％and
９４．８７％ofthecontrol,respectively(Table２)．InterＧ
estingly,evenafterbeingsubmergedinwaterfor３０d,
itsnetphotosyntheticrateat１６００μmol􀅰mＧ２􀅰sＧ１is
１４２．６１μmolCO２􀅰kgＧ１ DW􀅰sＧ１,being１１０．８７％ of
thecontrolsignificantlyhigherthanthecontrol．OverＧ
al,thenetphotosyntheticrateofB．procumbensin
waterdecreasesfirstlyandthenincreasesgradualy
withtheextensionofsubmersiontime(Table２)．
PnofHedwigiahamulosumafterimmersionin
Table１　PhotosyntheticlightＧresponsecurveequationsandphotosyntheticparametersoffourmossspecies
Mossspecies PhotosyntheticlightＧresponsecurveequations
MaximalPn
(μmolCO２􀅰
kgＧ１DW􀅰sＧ１)
LSP
(μmol􀅰mＧ２􀅰sＧ１)
LCP
(μmol􀅰mＧ２􀅰sＧ１)
Brachytheciumprocumbens Pn＝９６．１９􀅰PAR/(０．４９􀅰PAR＋１９５)－２４．４８,r＝１．００∗ １２２．５８±１．７３ １０１８．６０ ５６．７５
Hedwigiahamulosum Pn＝３８．２４􀅰PAR/(０．３１􀅰PAR＋１１５)－１．３２,r＝０．９９∗ ８７．２４±０．７１ ９４１．７６ ５．３０
Leucobryumglaucum Pn＝９．９１􀅰PAR/(０．１１􀅰PAR＋９１)－１．６０,r＝０．９８∗ ４１．１０±１．４５ ６７０．０３ ２０．５０
Hedwigiaciliata Pn＝３４０．２８􀅰PAR/(３．７４􀅰PAR＋９１)－７０．９４,r＝０．９７∗ １９．１０±０．２５ １１６６．００ ８５．００
　∗:atthe０．０１levelofsignificance．
Table２　NetphotosyntheticratesofBrachytheciumprocumbensaftersubmersion
inwaterfordifferentdays(μmolCO２􀅰kgＧ１DW􀅰sＧ１)
Immersed
days
Photosyntheticactiveradiation(μmol􀅰mＧ２􀅰sＧ１)
１２０ ２００ ４００ ６００ ８００ １２００ １４００ １６００
０ １８．３７±０．６７a ４１．０６±１．４２a ７２．５６±０．９０a ９２．３８±０．７１a １０６．４４±０．４６a １２６．７０±０．９９a １２９．５６±０．３５a １２８．７４±１．７３a
５ １７．０４±０．５４ac ３４．３１±１．１２b ６５．２４±０．９４b ８３．１１±０．９１b ９８．０３±１．１４b １２３．０４±１．１８a １２３．４７±１．７９b １２７．１２±０．７１a
１０ １１．５８±１．２１b ２７．５５±０．８７c ４５．８９±０．７８c ６３．８５±０．９９c ７４．９４±１．０５c ９７．１６±０．６０b １００．３９±０．５４c １００．６８±０．５９b
１５ １９．４６±２．９５a ２７．５７±０．５５c ４７．９７±１．２７c ６３．９１±０．９６c ７９．８３±１．９３d １１０．１５±２．８０c １０８．９８±２．９０e １１３．００±１．５６c
２０ １３．９９±１．１６bc ３０．５８±１．５７c ６２．１１±１．１６b ８０．１７±１．１１b ９１．８１±０．８６e １１７．２４±０．７９d １１９．３３±１．５２b １２２．１３±１．０５d
３０ １７．５２±０．２８ac ４７．４３±２．２７d ７１．４９±２．５０a ９７．８１±１．７２d １１９．３１±２．１３f １３３．７８±０．６１e １３８．９２±０．８０e １４２．６１±１．３８e
Significant
test
df＝２９
MSE＝１０．２４８
F＝４．２９２
P＝０．００６
df＝２９
MSE＝９．９２５
F＝３２．３９７
P＝０．０００
df＝２９
MSE＝９．６０４
F＝６８．８００
P＝０．０００
df＝２９
MSE＝６．１８９
F＝１６１．４６３
P＝０．０００
df＝２９
MSE＝９．６５４
F＝１４２．３２８
P＝０．０００
df＝２９
MSE＝９．６４４
F＝８７．７８５
P＝０．０００
df＝２９
MSE＝１２．４６３
F＝７７．６６０
P＝０．０００
df＝２９
MSE＝７．７６２
F＝１３２．８３７
P＝０．０００
　Note:Datawereaverageoffivereplications±SEMsandthesamesuperscriptletterwithinarowmeansnodifferenceatthe０．０１levelofsignificancebytheLSD
test．Thesamebelow．
water:Hypnumhamulosum,ifsubmergedinwater
for２０and３０d,itsnetphotosyntheticratesat５０
μmol􀅰mＧ２􀅰sＧ１are７．８０and７．３４μmolCO２ 􀅰kgＧ１
DW􀅰sＧ１,respectively,significantlylowerthantheconＧ
trol(１０．６０μmolCO２􀅰kgＧ１ DW􀅰sＧ１),whilethose
submergedfor５,１０,１５datthesamelightintensity
aresignificantlyhigherthanthatofthecontrol．OverＧ
al,thenetphotosyntheticrateofH．hamulosuminＧ
creasesfirstlyandthendecreasesgradualywiththe
extensionofsubmersiontime,butthenetphotosynＧ
theticratesofH．hamulosumat２００,３５０,５５０,７５０,
１０００,１１００μmol􀅰mＧ２􀅰sＧ１,evensubmergedinwater
for３０d,are９６．６３％,１０４．８２％,１０６．１０％,９５．５５％,
１０４．０５％ and９６．９６％ ofthecontrol,respectively．
Therefore,H．hamulosumisabletosurviveinwater
after３０Ｇdaysubmersion(Table３)．
００７ 广　西　植　物　　　 　　　　　　　　　　　　　　３５卷
Table３　NetphotosyntheticratesofHedwigiahamulosumaftersubmersion
inwaterfordifferentdays(μmolCO２􀅰kgＧ１DW􀅰sＧ１)
Immersed
days
Photosyntheticactiveradiation(μmol􀅰mＧ２􀅰sＧ１)
５０ １００ ２００ ３５０ ５５０ ７５０ １０００ １１００
０ １０．６０±０．９６a ２４．４３±１．３９a ３９．３０±１．３７ab ５４．１０±２．０４a ６７．００±１．２２a ８２．８９±０．９５a ８４．１１±１．８７ab ８９．０５±０．７１a
５ １１．５１±０．５５a ２７．０９±０．９８ab ４３．９７±１．０１ac ６３．５３±０．６６b ８３．４５±０．４３b ９９．２０±１．４０b １０６．６２±０．９２c １０６．８７±２．９２b
１０ １３．２１±０．８３b ２２．７４±１．２０ac ４０．１７±１．８０ab ５４．５５±０．５２a ７０．３７±０．５７cd ９９．３４±１．３４b １０２．６０±１．９７cd１０３．４６±０．７７b
１５ １２．５３±２．９３b ２８．９４±２．６６bde４６．８０±２．９７cd ６３．４５±２．４２b ７５．６６±１．９７e ８６．５０±０．９９c ９１．７２±０．４６e ９４．２９±０．２６c
２０ ７．８０±０．５４c ２５．３６±１．００ae ４１．０１±０．４４ae ５３．９３±０．４８a ６９．７２±０．４４adf ７７．１３±１．５９d ８２．４２±１．５５a ８３．０６±０．６３d
３０ ７．３７±０．５９c １９．７５±１．３８cf ３７．９７±０．９０be ５６．７１±０．５２a ７１．０９±１．０８cf ７９．２０±０．７４d ８７．５１±１．０７b ８６．３４±１．０９ad
Significant
test
df＝２９
MSE＝９．２８１
F＝３．１８８
P＝０．０２４
df＝２９
MSE＝１１．９０１
F＝４．４１４
P＝０．００５
df＝２９
MSE＝１３．２９５
F＝４．０１９
P＝０．００９
df＝２９
MSE＝９．３３８
F＝１１．２６４
P＝０．０００
df＝２９
MSE＝６．０４１
F＝２８．７１０
P＝０．０００
df＝２９
MSE＝７．２６５
F＝６５．４９３
P＝０．０００
df＝２９
MSE＝９．９５９
F＝５０．０９５
P＝０．０００
df＝２９
MSE＝９．３９８
F＝４８．７３３
P＝０．０００
Table４　NetphotosyntheticratesofLeucobryumglaucumaftersubmersion
inwaterfordifferentdays(μmolCO２􀅰kgＧ１DW􀅰sＧ１)
Immersed
days
Photosyntheticactiveradiation(μmol􀅰mＧ２􀅰sＧ１)
５０ １５０ ２５０ ３５０ ５００ ６５０ ８００ ９００
０ ５．１３±０．４４ab １０．４２±０．１６a １７．６０±１．０４a ２５．０９±０．３３a ３２．８０±０．８６a ４１．４９±１．１２a ４４．１５±１．４６a ４２．７０±１．３５a
５ １３．８７±０．４５c ２４．７３±１．４５b ３５．２７±０．３３b ４２．２８±０．７８b ４８．７８±０．８７b ５１．０３±０．９２b ５１．８３±０．５７b ５０．４５±０．７６b
１０ ６．８３±１．７２a １９．７１±１．４２c ３２．４２±０．９８c ３７．３６±０．６１c ４７．１９±０．６０b ４７．３９±０．６７c ４７．４７±０．５７c ４７．４４±０．９７c
１５ ６．８３±０．６４a １９．４８±０．８４c ２６．６９±０．４３d ３１．４８±０．５０d ３９．１９±０．３７c ４４．６９±１．０６d ４６．０７±０．６９ac ４６．２７±１．２３c
２０ ６．１４±０．７５a １８．６１±０．６６c ３２．５３±０．４６c ４０．４２±０．４９e ４７．１５±０．５３b ５２．４１±０．５１b ５７．１８±０．４６d ５７．６６±０．２１d
３０ ３．３７±０．８７b １７．６９±０．５０c ２７．５５±０．３１d ３３．３０±０．２６f ３８．６２±０．３６c ４２．４９±０．２２ad ４５．９９±０．４７ac ４６．７５±０．２２c
Significant
test
df＝２９
MSE＝４．２０８
F＝１５．３８２
P＝０．０００
df＝２９
MSE＝４．６０９
F＝２３．２４８
P＝０．０００
df＝２９
MSE＝２．１９３
F＝９１．３２０
P＝０．０００
df＝２９
MSE＝１．３６１
F＝１４８．１５９
P＝０．０００
df＝２９
MSE＝１．９９１
F＝１０１．９５６
P＝０．０００
df＝２９
MSE＝３．３１４
F＝３０．３９０
P＝０．０００
df＝２９
MSE＝３．０６８
F＝３８．５０３
P＝０．０００
df＝２９
MSE＝４．１０６
F＝３１．７６０
P＝０．０００
Table５　NetphotosyntheticratesofHedwigiaciliataaftersubmersion
inwaterfordifferentdays(μmolCO２􀅰kgＧ１DW􀅰sＧ１)
Immersed
days
Photosyntheticactiveradiation(μmol􀅰mＧ２􀅰sＧ１)
１００ １６０ ３００ ５００ ７００ ９００ １０００ １１００
０ ３．３８±０．４２a ７．６４±０．３９a １１．０２±０．４７a １４．３０±０．１３a １６．５６±０．１７a １９．０１±０．０９a １８．８２±０．２８a １９．１０±０．２５a
５ ２．９１±０．１０a ６．０８±０．１７b １０．１７±０．１５a １３．４０±０．１２b １６．６１±０．１１a １７．９３±０．１７b １８．３１±０．１８a １８．７０±０．２１a
１０ ０．８４±０．４３b ３．３５±０．２９c ６．７０±０．１６b ８．３２±０．２６c １１．６３±０．１８b １３．３１±０．１１c １３．７９±０．１２b １４．６１±０．１６b
１５ ND ２．８９±０．３４c ７．１０±０．４８b １１．９１±０．１８d １４．１６±０．１１c １６．１３±０．１０d １６．９６±０．０７c １７．８２±０．０７c
２０ ND ND ０．１７±０．０４c ０．４８±０．０４e ０．６３±０．０１d ０．８６±０．０２e ０．８２±０．２１d ０．９９±０．０２d
３０ ND ND ND ND ND ND ND ND
Significant
test
df＝１４
MSE＝０．６２２
F＝１４．６９７
P＝０．００１
df＝１９
MSE＝０．４７４
F＝５３．７１７
P＝０．０００
df＝２４
MSE＝０．４９８
F＝１８２．８６０
P＝０．０００
df＝２４
MSE＝０．１３２
F＝１１９９．０６８
P＝０．０００
df＝２４
MSE＝０．０８６
F＝２５６５．２０８
P＝０．０００
df＝２４
MSE＝０．０６２
F＝４４０１．１７２
P＝０．０００
df＝２４
MSE＝０．１７２
F＝１６２５．６７７
P＝０．０００
df＝２４
MSE＝０．１３４
F＝２１６３．０６１
P＝０．０００
　　PnofLeucobryumglaucumafterimmersionin
water:ThenetphotosyntheticratesofL．glaucum,
exceptthatsubmergedinwaterat５０μmol􀅰mＧ２􀅰sＧ１
for３０d,aresignificantlyhigherthanthecontrol(TaＧ
ble４)．Aftersubmergedinwaterfor５,１０,１５,２０and
３０d,thenetphotosyntheticratesofL．glaucumat９００
μmol􀅰mＧ２ 􀅰sＧ１are５０．４５,４７．４４,４６．２７,５７．６６,and
４６．７５μmolCO２ 􀅰kgＧ１ DW􀅰sＧ１,respectively,being
１１８．１６％,１１１．１１％,１０８．３７％,１３５．０３％and１０９．４９％
ofthecontrol,respectively,indicatingthatL．glaucumis
alsoabletosurviveinwater,atleastfor３０d(Table４)．
PnofHedwigiaciliataafterimmersioninwaＧ
ter:Afterbeingsubmergedinwaterfor１０－２０d,the
netphotosyntheticrateofH．ciliataslightlyincreased
firstly,thendecreasedsharply,nonetphotosynthetic
rateofH．ciliatawasdetectedaftersubmergedinwaＧ
１０７５期　　　　　　　申琳等:基于光合参数探讨四种藓类作水族箱植物的应用潜力
terfor３０days(Table５),indicatingthatHedwigia
ciliataisnotadaptedtowaterenvironmentover１０d．
３　Discussion
TheLCPofshade herbsarelowerthan ２０
μmol􀅰mＧ２􀅰sＧ１,andtheirLSParefrom５００to１０００
μmol􀅰mＧ２􀅰sＧ１orlower(Niuetal．,２００４)．TheLCP
ofH．hamulosumandLeucobryumglaucumare５．３０
and２０．５０μmol􀅰mＧ２􀅰sＧ１,respectively,andtheirLSP
are９４１．８０and６７０．００μmol􀅰mＧ２􀅰sＧ１,respectively,
indicatingthatbotharetypicalshadeplants．Compared
withL．glaucum,Hedwigiahamulosum haswider
light adaptation range under natural condition;
BrachytheciumprocumbensandHedwigiaciliatabeＧ
longtosunnyplants,whichisrevealedbytheirhigher
LCPfrom５６．７５to８５．００μmol􀅰mＧ２􀅰sＧ１,andalso
higherlight saturation points from １０１８．６０ to
１１６６．００μmol􀅰mＧ２􀅰sＧ１．ComparedwithBrachyＧtheＧ
ciumprocumbens,HedwigiaciliataisatypicalhelioＧ
phyte,similartootherterrestrialsunnyherbsinitsadＧ
aptationtolight．
TheLCPandLSPofthefourmossspeciesare
relatedtotheirnaturalhabitats．ThesamplesofHedＧ
wigiaciliata weretakenfromrocksurfaceinopen
field．H．ciliata,anextremelydroughtＧtolerantmoss
species,oftendistributesonsurfaceofopenrockswith
stronglight;Brachytheciumprocumbensoccasionaly
growsongrasslands,hilsinsunnyandopenhabitats,
butalsoonfloorandbouldersunderforest(Huetal．,
２００５)．ThesamplesofB．procumbensweretakenfrom
roadsideofasparseforestinXuhuiCampusofShangＧ
haiNormalUniversity．ItshigherLCPandlowerlight
saturationpointindicatingthatB．procumbensisneiＧ
theratypicalshadeplant,noratypicalsunnyplant．
ThesamplesofHedwigiahamulosum weretaken
fromapatchofshrubbyroadside(evelvationca．１１８０
m)ofJinhuaMountain,thehabitatisshadyandoften
misty,thephotosyntheticparametersofH．hamuloＧ
sumreflectsthehabitatconditionstosomeextent．
FourmossspeciesvarymuchintheirPn．Among
them,BrachytheciumprocumbenshasthehighestvalＧ
ue,being１２２．５７５μmolCO２􀅰kgＧ１DW􀅰sＧ１,whileH．
ciliatathelowest,being１９．１０μmolCO２􀅰kgＧ１DW􀅰
sＧ１,theformerhas６．４２timeshigherPnofthatofthe
latter．Theirphotosyntheticparametersarerelatedto
theirmorphologicaltraits．ThereasonforH．ciliata
haslowerPncouldbeexplainedasfolows:(１)HedＧ
wigiaciliataaregreygreenishwithlowerchlorophyl
contentcomparedwithBrachytheciumprocumbens,its
leafhyalinetipsarenotaphotosynthetictissue,acＧ
countingforalargepartofleaf;(２)itsleavesdensely
coveredwithmanysharptransparentpapilaeonboth
sides,which mayafecttheabsorptionoflight;(３)
Hedwigiaciliatahasprocumbentandstoutstems,the
photosyntheticcapacityofthispartwilcertainlybe
weakerthantheleaves．
ThePnofLeucobryumglaucumis４１．１０μmol
CO２􀅰kgＧ１ DW􀅰sＧ１,higherthanthatofHedwigia
ciliata,but significantly lower than those of
BrachytheciumprocumbensandHedwigiahamuloＧ
sum,whichisrelatedtoitsgametophytefeatures．The
leavesofLeucobryumglaucum havesmoothsurface
withoutpapiliaeandhyalinetipslikeHedwigiaciliaＧ
ta,buttheyhaveflatandwidecostae,andonlyone
layerofgreencelswithphotosyntheticfunction,and
thelargercelsonbothsidesofthegreencellayerare
colorless,withoutphotosyntheticpigments．
MostpreviousreportsaboutbryophytephotosynＧ
thesistookleafareaasunit,sowecannotcompareour
photosyntheticdatawiththoseofpreviouswork．DelＧ
toroetal．(１９９９)reportedthatthePnofLeucodon
sciuroidesinMediterraneanwas８－１０mgCO２􀅰gＧ１
DW􀅰hＧ１ at１５ ℃,whichisequalto５０－６０μmol
CO２􀅰kgＧ１ DW􀅰sＧ１,andclosetothatofLeucobryum
glaucuminthepresentwork．Convey(１９９４)reported
thatthePnofBartramiapatensandothertwelve
mossspeciesintheAntarcticvariedfrom０．８７９to
０．１３４mgC􀅰gＧ１DW􀅰hＧ１．ifconvertedintothesame
unit,thePnofBrachytheciumprocumbensis０􀆰８２－
４􀆰５０mgC􀅰gＧ１DW􀅰hＧ１．ConsideringtheharshAntＧ
arcticenvironment,thePnofthebryophytesmeasured
byConveyarenaturalylowerthanthoseofthepresＧ
entwork．Therefore,thephotosyntheticdataofthe
fourmossspeciesfalintothereasonablescope．Ueno
etal．(２００６)reportedthenetphotosyntheticrateof
２０７ 广　西　植　物　　　 　　　　　　　　　　　　　　３５卷
Calliergongiganteum (asemiＧaquaticmossspecies)
as１􀆰２－１􀆰６mgCO２􀅰gＧ１􀅰hＧ１,about１１３－１５０μmol
CO２ 􀅰kgＧ１ DW 􀅰sＧ１,whichissimilartothatof
Brachytheciumprocumbens．Liuetal．(２００１)found
thatthePnofPlagiomiumacutumandP．maximovＧ
icziinsummeris１２５．６７and９４．６３μmolCO２􀅰kgＧ１
DW􀅰sＧ１,whichshowsthatPlagiomiumacutumand
Brachytheciumprocumbens,PlagiomiummaximovicＧ
ziandHedwigiahamulosumhavesimilarphotosynＧ
theticcapacity,respectively．
Aftersubmergedfordiferenttimes,thePnof
Brachytheciumprocumbensincreasesfirstlyandthen
decreasesgradualywiththeextensionofsubmersion
inwateratalPARintensitiesexceptfor１２０μmol􀅰
mＧ２􀅰sＧ１．ThephotosynthesisofBrachytheciumprocＧ
umbens,ifsubmergedforashorttime,wasinhibited
becauseofwaterstressinrelationtodeficiencyofoxyＧ
genandCO２．Withtheextensionofsubmergedtime,
B．procumbensisgradualyadaptedtowaterenvironＧ
mentanditsphotosyntheticcapacityrecovered,andeＧ
venexceededthecontrol．Duringthesubmersion,the
gametophytesofB．procumbenskeptgreenalthe
time,indicatingthatB．procumbensisabletosurvive
welinaquaticenvironment．Inthefield,B．procumＧ
bensoftendistributesonstonesonedgeofsome
streams,sometimessubmergesinwaterduringrainy
season．
ThenetphotosyntheticrateofLeucobryumglauＧ
cumincreasedfirstlyandthendecreasedgradualywith
theextensionofsubmersion,itsPnreachedapeakafＧ
terbeingsubmergedfor２０d,indicatingthatLeucobryＧ
umglaucumisabletolivewelinwaterforashort
time．ThegametophyteofL．glaucumissomewhat
similartothatofSphagnumsp．,saySphagnumpalＧ
ustre,thelatterisabletolivewelinswampsandwet
environments．Therefore,L．glaucumhasmorphologiＧ
calandstructuralgroundtoadapttoaquaticenvironＧ
menttoacertaindegree．Whensubmergedinwater,
Hedwigia hamulosum and Leucobryum glaucum
shareasimilarchangepatternintheirPn．TheirPn
increasedfirstlyandthendecreasedgradualywiththe
extensionofsubmergedtime,buthigherornotsignifiＧ
cantlylowerthantheircontrol,respectively,evenafter
３０Ｇdaysubmersioninwater,showingtheirgoodadaptＧ
abilitytowaterenvironment．
HedwigiaciliataisadroughtＧtolerantsaxicolous
mossspecies．ThePnofH．ciliatadecreasedsharply
withtheextensionofsubmergedtime,indicatingthat
H．ciliataisnotadaptedtoaquaticenvironment．
４　Conclusions
Overal,Brachytheciumprocumbens,LeucobryＧ
umglaucumandHedwigiahamulosumareadaptedto
aquaticenvironmentstosomeextent,andtheyseems
tobepotentialaquariumplants．Itshouldbenotedthat
thisstudyhasmeasuredthePnofonlyfourmosses
aftertheirbeingsubmerged,ifweextendourpresent
workwithmoreotherterrestrialsandwetmossspeＧ
cies,wemaybefindmorespeciesadaptedtowaterenＧ
vironments,andthusprovidemorenewpotentialaＧ
quariumplants．
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３０７５期　　　　　　　申琳等:基于光合参数探讨四种藓类作水族箱植物的应用潜力