BulletinofGlaciologicalResearch22（2005）6974∂9◎JapaneseSocietyofSnowandIce

Determinationol＝aminoacidsinicesamples
FumioNAKAZAWAl．KeiichiOHTA2．NaomiHARADA3，KojiFUJITA4

andMasaYOShiNAKAWO5

1GraduateSchoolofScience，NagoyaUniversity，Furo−Cho，Chikusa−ku，Nagoya464−8601，Japan

2DepartmentofEcosystemStudies，SchoolofEnvironmentalScience，UniversityofShigaPrefecture，2500Hassaka−Cho，Hikone，
ShigaPref．5228533，Japan

3JapanAgencyforMarine−EarthScienceandTechnology，215Natsushima−Cho，Yokosuka−City，Kanagawa2360061，Japan
4GraduateSchoolofEnvironmentalStudies，NagoyaUniversity，Furo−Cho，Chikusa−ku，Nagoya4648601，Japan

5ResearchInstituteforHumanityandNature，335Takashima−Cho，Marutamachi−doriKawaramachinishi−iru，KamlgyO−ku，Ky
6020878，Japan（ReceivedOctober4，2004；Revised manuscriptreceivedNovember30，2004）

Abstract
This studyinvestigates amino acid concentrations and compositionsinice as a preliminary

approachforapplylngaminoacidanalysistoicecorestudy．Fortheaminoacidanalysis，0．425toO．750
kgicesamplesare used．Theicesfrom Antarcticahave16．8and18．3nmolkg10faminoacidsand

theseconcentrationsareclose tolaboratory blanklevels，Whiletheicefrom theChongceIceCapin

Chinahas160nmolkg10faminoacids．Therefore，theanalysisforAntarcticiceisestimatedtoneed
morethanseveralkgofice．Incontrast，theanalysisfromtheChongceIceCapshouldbereducibleto

Only severalhundred grams ofice because the concentrationsin this cap are higher than thosein

Antarctica．Further，amino acid sources are presentin glacier surroundings of temperate reglOnS，WhereaspolarreglOnSarefarfromsources．Serine，aSparticacidandalaninearethemaJOrCOmpOnentS
inthesamples．However，theaminoacidcompositionsshowvariationwitheachsample．

1．lntroduction

Seem tOOrlglnatemainlyfrom microorganisms，SuChas snow algae and bacteria，that breed on glacierSurfacesin summer season（Kohshima，1987，1989；
Ling and Seppelt，1990，1993；Yoshimura etal．，1997；
Takeuchietal．，2001），plantproducts，SuChaspollen，inglacier surroundings（Koerneretalり1988；Bourgeois，
1990，2000；Liuetal．，1998；Reeseetal．，2003；Nakazawa
eventsincludingbiogenicemission，biomassburning， etal．，2004）andthebacteriainprecipitation（Herlihyet
anthropogenic emission and voIcanic eruption（e．＆， al．，1987；Sattleretal．，2001；Maceetal．，2003）．Thus，theHammer，1977；Hammer etalり1980；Legrand andamino acids may have good environmentalinforma−
tioninicecorestudy．Sincethesebiologicalactivities
Angelis，1996；01ivier etal．，2003）．However，there are

Ionic species，SuCh asinorganicions andlight
Carboxylicacids，inicecoreshaveprovidedimportant
information on pastclimatic and environmentalcon−
ditions．The chemicalspecies yielddataon historical

Wheninterpretationis difncult．Forexample，natural

increasein summer season and decreasein winter，amino acid−richlayersin anice core mayindicate

fires，increase concentrations of chloride，nitrate andSulfateionsintheatmosphere．Inaddition，anthropo−

ineachsummerlayermaybeusefulasagoodmarker
OfmeteorologlCalconditionssuchastemperatureandsome other candidates for the source tracers in cases
events，SuCh as voIcanic eruptions andlarge forest

Summerlayers．Inaddition，aminoacidconcentrations

genic effects such as biomass burning and fossil fuelCOmbustioncanalsoincreasetheatmosphericconcen−
trations of these constituents．

glacierice．Therefore，aSapreliminaryapproachtoice

entlyoriginatefromandareabundantinlivingorgan−amino acid concentrationsinice from the Antarctic

Incontrast，aminoacidsthatbuildproteinsappar−

SOlar radiation that affect these activities．However，

no previous study has reported the amino acids inCOre reSearCh，this study aims to survey thelevelof

isms，forexample，apprOXimately50％of the human icesheetandChongceIceCapincentralAsia，andto
body sdryweightisprotein（McMurryandCastellion， COmparetheircompositions．
1999）．Moreover，aminoacidscontainedinglaciericesBu11etin ofGlaciologlCalResearch

7P

Tablel．Detectionsystemforhigh−performanceliquidchromatography（HPLC）．
HPLC

Shimadzu LC−10AD

Integrator
ColumnShimadzuchromatopacGR7Aeplus
GLScienceIntersilODS−3（150mm L，4．6mmID，5〟mparticlesize）

Detector

Guard column
Column temperature
EluentFlow−rate

FluorescencedetectorRF−10AXL（350¶460nm）

GLScienceIntersilODS−3（50mmL，4．6mmID，5pmparticlesize）
400c
El：5mMNa2HPO4・12H20／5mMNaH2PO4・2H20（pH7．05）
E2：El／CH3CN（2：1，V／v）
E3：CH3CN／Distilled water（HPLCgrade）（4：1，V／v）
1．OmL minl

2．SampllngSitesandsampIepreparation
Ice core sampling was conducted at two distinct

sites；AsukacampinEastAntarctica（71031′s，2408′E；
930m a．s．l．）and ChongceIceCapin theWestKunlun
Mountains，China（35014′N，81007′E，6327m a．s．1．）．Ice

blocks were taken from the surface ofanicebergoff

Table2．Working gradient conditions used for aminoacid determination．
Time（mim）

EI

E2

E3

Programl Program2
0．01

0．01

4．0

4．0

8．0

8．0

16．0

16．0

5

0

5

0

0
7
1

0

2

5

2

0

3

5

4

0

5

5

6

0

7

0

7

0

8

0

0

0

0
0
0
︵
U

Syowa Station（69000′s，39035′E）in the Antarctic
18，0
Ocean．One section was collected from eachice core；
25，0
fromtheAsukacore，theincrementbetweenlOO．8and
30．0
lOl．3mdepth，WhilefromtheChongcecore，theincre− 42．0
mentbetween2．8and3．Om depthwereused．Forthe 42．01
aminoacidanalysis，0．750，0．500andO．425kgiceblocks
45．0

32．0

O

44．0

O

44．01

O

51．0

53．0

51．01

53．01

52．0

54．0

84．0

90．0

84．01

90．01

Were uSed for the Asukaice，theicebergice and the
Chongceice，reSpeCtively．

TopreventcontaminationbylaboratorycontroIs，precleaned glasswares，Whichinclude filters，Wereusedin al11aboratory work．The glasswares werepreparedbywashinglnalkalinedetergent，placingln
IN HCI overnight followed by rinsing with organic‑
free water obtained with a Milli−Q system and thenheatinginafurnaceat4500cfor3to4h．Additionally，

toeliminatecontaminationofeachsample，aboutlcm
Ofthesurfacewasscrapedoffwithacleanknifeina

COld room，and thenthesamplewasmeltedinabea−
ker．Thefirst meltedice used to wash the surface oftheice samples was discarded．This procedure was

18．0
27．0

O
O
O
O

0

O

0

O
O

47．0

84．02（stop）90．02（StOp）

O0
0

The programlwas used for the particulate anddissolved fractionsofthe blanksample，the particulate
fraction of the Asuka sample，and the particulate
fraction of the Chongce sample．The program2wasused for the dissolved fraction of the Asuka sample，
the particulate and dissolved fractions of the iceberg
Sample，and the dissoIved fraction of the Chongce
Sample．

repeated3to5times，depending on the sample．The
meltedicediscardedinthiswaycametoO．145toO．110 acid content of theices．
kg．After theice was completely melted，the melt
Alaboratory proceduralblank were also ana−
WaterWaSimmediatelyfi1tratedthrough47mmGF／F
1yzed．Forparticulateaminoacids，aneWGF／Fglass
glassfiberfilterswithaporesizeofO．7／Jm．
fiberfi1ter（diameter：47mm；pOreSize：0．7FLm）washy−

Subsequently，the samples were separatedinto

drolyzedandthefollowingprocedurewasthesameas

twoportions，afi1tersampleandfi1tratesample．Thesethatforsamples．In contrast，fordissoIvedaminoacT
6sub−Samples（for amino acid analysis）were hydro−
ids，4−mloftheorganic−freewaterwasconcentratedin
1yzed with6N HClatlO50c for22h．The HCIsuper−anaskundervacuum，and passed throughthecation
natantwasthentransferredtoaflaskanddriedunder

exchange resin．The fo1lowing procedure was theizedwitho−phthalaldehydeandthenanalyzedbygraT

ent conditions used for determination of amino acidaresummarizedinTableland2respectively．Twelve

VaCuum．Afterdrying，thedried residue wasdesalted same as mentioned above．
usingcationexchangeresin（Dowex50W−Ⅹ8），derivat−
AdetectionsystemforHPLCandworkinggradi−
dient reversed−phase high−Performanceliquid chro−
matography（HPLC）with UVfluorescence detectionin order to determine the totalhydrolyzable amino

amino acids were measuredin this study（Table3）．

Theyieldforthepurificationofaminoacidswas95toNakazawaetal．

7ノ

Table3．Concentrationofaminoacidsintheice（nmolkg1）．Abbreviation：Glu（glutamicacid），Ala（alanine），Val（valine），
Gly（glycine），Ile（isoleucine），Ser（serine），Thr（threonine），Arg（arginine），Asp（aspartic acid），Met（methionine），Phe（phenylalanine），Tyr（tyrosine）．
Sample Type sample Glu Ala Val Gly Ile Ser Thr Arg Asp Met Phe Tyr Total
Blank r1

：i

．2

Asuka r，lceberg Particulatemetter O．4 1．2
DissoIved matter l，8 1．1 0．7
Chongce Particulatemetter 30 24 15
DissoIved matter O．5

0．9

0．3
0．5

0．4 1．2 0．8 1．O n．i． 0．9 n．i． n．i． 0・
1．4 1．9 1，5
0．2 l．4 1．2 n．i． 0．2 11．！）

2．7 7．9 20
1．7 0A n．i．

17
0．9

5．5 200．9 0．6

3．3
0．9

2．9 3・0 152
0．5 n．i．

0．2

7．5

toidentify thepeakofaminoacidbecauseamountswerebelow
Then．i．is notidentified ，meaningthatitwasimpossible
thedetectionlimitorinseparablefromadjacentpeaks．

100％（Harada，1991）．The precisions（RSD，n＝2）were
13tol％except for valine（Val），isoleucine（Ile），me−
thionine（Met），Phenylalanine（Phe）andtyrosine（Tyr）．

TheprecisionsofthefiveaminoacidswerenotdeterL
mined．

acid concentrationsin Antarcticice should begener−
allylow．

TheChongcesampleshowsthatGlu，Ala，SerandAsparethemaJOrCOmpOnentSOftheparticulatefrac−

3．Results and discussion

tion and comprise62％of the total．In contrast，Val，
Ala，Ser，ThrandAsparethemaJOrCOmPOnentSOfthe
dissoIved fraction and are71％ofthe total．The total

Totalconcentrationsofparticulateand dissoIved
amino acidsin theice samples from the three sites

Samples，SpeCifically，13timesthatoftheAsukaand24

range between16．8to160nmolkg1．Table31iststhe

concentrationsoftwelveaminoacidsintheblankand
3ice samples．The blank values are not subtracted
from the sample values because we did not measureamino acid concentrations in the organic‑free waterused fortheblanksampleand theblank values were

COnCentrationin theparticulatein theChongceiceis
muchhigherthanthoseoftheAsukaand theiceberg

times that of theiceberg，eVen though the dissolved
matterintheicesfromthethreesiteshavesimilarand
blanklevels．

The differencein particulate amino acid concen−
trations between the Antarctica and Chongce icesseemstobeattributabletothedifferenceinthesource

Closetothedetectionlimitsof2to3pmol．Therefore， Strengthin two reglOnS．Theice samplesinlower

it would be safe here to use the blank values asreferences．
The totalconcentrationsin the Asuka core andiceberg samples are at similarlevels．Moreover，theconcentrationofeachaminoacidis at the blanklevel
inboththeparticulateanddissoIvedfractions．Inthe

Asuka sample，Serine（Ser），Val，glutamic acid（Glu）
and alanine（Ala）are the major componentsin theparticulate fraction，and constitute67％of the totalaminoacids．Ser，aSParticacid（Asp），Tyrand Alaare

the maJOr COmpOnentSin the dissoIved fraction，andlatitudinalicecapsandglaciers（betweenabout600N
and600s）typically contain higherconcentrationsof

nutrients，SuCh as water−SOluble chemicalspecies，
than thatfrom thepolarice sheets（e．g．，Wagenbach，
1989；Clausen and Langway，1989；Lyons etal．，1991）．

The higher concentrations play the more importantrolein the breeding ofsnow algae．Moreover，On glaT

Ciersurfacesinlowerlatitudes，SOmeSurfacemelting
OCCurSinthesummerseason，andsnowalgaegrowin

the meltwater（Pollock，1970；Yoshimura etal．，2000）．

However，the surface of polarice sheetsis virtually
lifelessbecauseofthelackofmeltwater，eVenthough
COnStitute69％ofthetotal．Intheicebergsample，Ala，Ile，threonine（Thr）andAsparethemajorcomponents Viablemicrobialcellsandsporesarepresent（Catranis

in the particulate fraction，and constitute67％ofthe
total．Ser，Glu，Thr，Ileand AsparethemaJOrCOmpO−
nentsin the dissoIved fraction，and are67％of thetotal．The Asukaicesectionisestimated to have been
hundredsofyearsoldbasedonanaverageaccumula−tion rate of O．19m a10fice between1988and1991（Azumaetal．，1994）．In addition，theice from theice−berg was much older than the Asukaice becauseit
hadoncebeenapartoftheAntarcticicesheet．Aminoacidsseemtobewe11preservedinglaciericebecause
Ofitslow temperature．Therefore thelevelof amino

and Starmar，1991）．Thus，thelocalgrowth of snow

algaemaycausethe higherconcentrationofparticu−
1ate amino acidsin Chongceice compared to thatin
Antarcticice．

Inaddition，thewindTblown−particulateaminoac−
idssuchasin pollen，may alsocontribute totheCho−

ngceIce Cap．Ice cores fromlowerlatitudeice caps
and glaciers that are typica11y within a few tens ofkilometersfromvegetationsourcescontainmorethanlOOOgrainsL10fpo11en（Ambachetal．，1966；Haeberlietal．，1983；Liuetal．，1998，Reeseetal．，2003；Nakazawa

BulletinofGlaciologicalResearch

72

（b）

（d）

Fig・1・Aminoacid compositioninicesamples．（a）ParticulateaminoacidsintheAntarcticAsukacoreatadepthoflOO．8
to101．3m；（b）DissoIved amino acidsin the Antarctic Asukacore；（c）Particulateaminoacidsin theAntarcticiceberg；（d）DissoIved aminoacidsin the Antarcticiceberg；（e）Particulate aminoacidsin the ChineseChongceice atadepth
Of2．8to3．Om；（f）DissoIved amino acidsin the Chinese Chongceice．Abbreviation：Glu（glutamic acid），Ala（alanine），
Val（valine），Gly（glycine），Ile（isoleucine），Ser（serine），Thr（threonine），Arg（arginine），Asp（aspartic acid），Met（methionine），Phe（phenylalanine），Tyr（tyrosine）．

etal．，2004），Whereas the pollen of polarice sheets blanklevel．Thisfactseemstobeduetofewsourcesof

mostly originates from sources hundreds to thou−amino acidsin situ andin the surroundings．Itis

Sandsofkilometersaway．Thus，thepollenconcentra− therefore concluded that more thanseveralkgoficetioniniceistypica11yonlylOtolOOgrainsLl（McAndrews， SamPles would be needed for analysisin ordertobe
1984；Short and Holdsworth，1985；Koerner andWellabovetheblanklevel．However，icecoresamples
Bourgeois，1988；Bourgeois，1990，2000）．Judging fromare typica11ylimited，anditis therefore essentialto

theabove，theicefromlowerlatitudinalicecapsandglaciers can be expected to contain enough particu−

SeCure enOughice volume forice core studyin thepolaricesheetsofsuchasAntarctica．
1ateaminoacidsforanalysis．Thus，thesamplevolume
Ser，Asp and Alaare the maJOrCOmpOnentSin5
may be reduciable to about several hundred grams
0utOfthe6subTSamples（Fig．1）．Howevertheamino
COnSideringthelaboratoryproceduralblankiftheiceacid compositions show variation with each sample．
fromlowerlatitudeicecapsandglaciershasaconcen−
The three amino acids were also found to be maJOr
trationsimilartothatoftheChongceicesample．
COmpOundsin soilorganic matter，pOllen，marine al−
In contrast，the variousice from the polarice
gae and marine bacteria（e．g，Auclair andJamieson，
Sheets may be difncult to analyze usingonly several
1948；Bieberdorfetal．，1961；Gupta and Reuszer，1967；
hundred grams ofice samples．This study used theSimon and Azam，1989；Cowie and Hedges，1992；CaoO．750and O．500kgicesamplesfortheanalysisofAnt− and Xiang，1995；Senwo and Tabatabai，1998），al−arcticices，thereby showing that concentrations ofbothparticulateanddissoIved aminoacidsareatthethough the amino acid composition differs by type．
Theexactcauseofthe differencesin amino acid com−Nakazawaetal，

7プ

POSitionintheiceisnotyetknown．Inparticular，the Azuma，N．，Goto−Azuma，K．andNakawo，M．（1994）：Photogra−

metric analysis of alOOmice core from Asuka Camp，East Antarctica：Preliminary results．Bulletin of Glacicr
Research，12，6975，
amino acids．Thus，further researchis required to Bieberdorf，F．W．，Gross，A．L．and Weichlein，R．（1961）：Frceaminoacidscontentofpollen．AnnalsofAllergy，19，8（う9ascertain whether these differences are great enoughaminoacidcompositionin theparticulatefractionof
the Chongce sample may relate to some origin of

to have practicalsignincance．

876．

Bourgeois，］．C．（1990）：Seasonaland annualvariationofpoIL
len contentin the snow of a Canadian high Arcticice
Cap．Boreas，19，313−322．
4．Conclusion
Bourgeois，J，C．（2000）：Seasonalandinterannualpollen variL
abilityln SnOWlayers of arcticice caps．Review of Pa−
This study investigates the concentrations and1aeobotanyandPalynology，108，1736．
COmpOSitionsofaminoacidsinicesamplesfromthree
Cao，S．andXiang，B．（1995）：Acomparativestudyonprotein
content and amino acid composition of eight species ofSites．Weconcludethatalargeicesample（morethan
Unicellularmarinefeedalgae．ActaOceanologicaSinica，
SeVeralkg）would beneededforanalysisofAntarctic
14，141146．
ice，SuChastheAsukacoreandiceberg．Consequently， Catranis，C，and Starmer，W．T．（1991）：Microorganisms en−
itis essentialto secure enough sample volume fortrappedin glacialice．AntarcticJournalof the UnitedStates，26，234236．
COntinuousanalysisinicecorestudies．Incontrast，We
Clausen，H．B．and Langway，Jr．，C．C．」．（1989）：Theionic de−
COnCludethattheamountofseveralhundredgramsof
positsinpolaricecores．In TheEnvironmentalRecordln
iceis needed for amino acid analysIS Of thelower
Glaciers andIce Sheets．（H．Oeschger and C．C．］．Lang−latitudinalicecapsandglaciers，1iketheChongceIce
Way，Jr．Ed．），pp．225247，JohnWiley＆Sons，NewYork．
Cowie，G．L．and Hedges，］．I．（1992）：Sources and reactivities
Capof the West Kunlun Mountains，due to theshortofaminoacidsin acoastalmarine environment，Limnol−

distancefromaminoacidsources，SuChassnowalgae，OgyandOceanography，37，703724．
bacteriaandpollen．Therefore，aminoacidanalysISin Gupta，U．C．andReuszer，H．W．（1967）：Effectofplantspecies

ice cores fromlowerlatitude reglOnS SeemS tO beeasierthanthatinthosefromthepolarreglOnS．
On the other hand，the cause of the different
amino acid compositions among the samples is anon the amino acid content and nitrification of soilor−ganicmatter．SoilScience，106（6），395−400．
Haeberli，W．，Schotterer，Uり Wagenbach，D．，Haeberli−
Schwitter，H，and Bortenschlager，S．（1983）：Accumula−tion characteristics on a cold，high−alplnefirn saddle
from a snowLpit study on Colle GnifettiMonte Rosa，unsettledquestion．Theaminoacidcomposition，eSpe−
SwissAlps．JournalofGlaciology，29，260271．
Cially thatin the particulate fraction oftheChongce
Hammer，C．U．（1977）：Past voIcanism revealed byGreenland
Sample，mayrelatetothesourcesofaminoacids，andicesheetimpurities．Nature，270，482−486．
allowustomoreeasilyprovidefurtherinformationto
Hammer，C．U．，Clausion，H．B．and Dansgaard，W．（1980）：Gre−enlandice sheet evidence of post−glacialvoIcanism andinterpret the past climate and environmentalcondi−
itsclimateimpact．Nature，288，230−235．
tions．Thus，thereisroomforfurtherinvestigationon
Harada，N．（1991）：Study on chronology ofmarinesediments
thispoint．
by the racemization reaction of amino acidsin plank−
tonic foraminifera（inJapanese）．M．S．thesis，98pp．，NaL
goyaUniversity，
AcknowLedgments
Herlihy，］．LリGa1loway，J．N．and Mills，A．L（1987）：Bacterial
utilization of formic and acetic acidin rain water．At−
WewouldliketothankJARE30forprovidingthemosphericEnvironment，21（11），23972402．
Koerner，R．MリBourgeois，J．C．and Fisher，D．A．（1988）：Pollen
ice samples from Antarcticaand theiceberg．Thanks
analysis and discussion of time−SCalesin Canadianice
are also due to N．Azuma of Nagaoka University ofcores．AnnalsofGlaciology，10，8591．
Technology for providing the density data from theKohshima，S．（1987）：Formation of dirtlayers and surface
Asukaicecore，andtoananonymousreviewerforhis
dustbymicro−PlantgrowthinYala（Dakpatsen）Glacier，
NepalHimalayas．BulletinofGlacierResearch，5，63−68，
helpfulcomments toimprove the manuscript．The
Kohshima，S．（1989）：Glaciologicalimportance of microL
field researchandanalysisweresupported byGrant−
Organismsin the surface mud−1ike materials and dirt
in−Aid for Scientific Research（Project Nos．12878087
layerparticlesoftheChongceIceCapandGozhaGlacit！r，and13373006）andthe21stCenturyCOEProgram（No．
West Kunlun Mountain，China．Bulletin of Glacier Re−
SearCh，7，5966，
G−4，Nagoya University）from the MinistryofEduca−
Legrand，M．and Angelis，M．D．（1996）：Lightcarboxylic acids
tion，Culture，Sports，Science and Technology ofinGreenlandice：Arecordofpastforestfiresandvegeta−
Japan．
tion emissions from the borealzone，Journalof Geo−physicalResearch，101（D2），4129−4145．
Ling，H．U．andSeppelt，R．D．（1990）：SnowalgaeoftheWind−
References
millIslands，COntinentalAntarctica．Mesotaenium beγg−
greni（Zygnematales，Chlorophyta）thealgaofgreysnow．
Ambach，W．，Bortenschlager，S．and Eisner，H．（1966）：Pollen−
Antarctic Science，2，143148．
analysis investigation of a 20m firn pit on the Kessel‑
Ling，H．U．andSeppelt，R．D．（1993）：SnowalgaeoftheWind−
Wandferner（OtzalAIps）．Journalof Glaciology，6，233
millIslands，COntinentalAntarctica．2．Chloromonasrub−
236．
roleosasp．nov．（Volvocales，Chlorophyta），analgaofrcd
Auclair，J．L．andJamieson，C．A．（1948）：A qualitativeanaly−
SnOW．EuropeanJournalPhycology，28，7784．
Sis of amino acidsin pollen collected by bees．Science，
Liu，K．BリYao，Z．andThompson，L．G．（1998）：Apollenrecord
108，357358．
Of Holocene climatic changes from the Dundeice cap，

74

Bulletin ofGlaciologicalResearch

Qinghai−TibetanPlateau．Geology，26，135−138．
persaland deposition on theIceCap ofMt．Parinacota，
Lyons，W．B．，Wake，C．，Mayeeski，P．A．（1991）：Chemistry of SOuthwestern Bolivia．Arctic，Antarctic，and Alpine Re−
SnOW from high altitude，mid／lowlatitude glaciers．In
SearCh，35（4），469474．
5eαざ0乃dJ5乃0比少αC々∫，MTOノ1SJSgγgeS，mJ．C2＆（T．D．
Sattler，B．，Puxbaum，H．andPsenner，R．（2001）：Bacterialgro−
Davies，M．Tranter and H．G．Jones，Ed．），pp．359−383，
Wthinsupercooledclouddroplets．GeophysicalResearch
Springer−Verlag，Berlin Heidelberg．
Letters，28（2），239−242．
Mace，K．A．，Kubilay，N．and Duce，R．A．（2003）：Organic nitro− Senwo，Z．N．andTabatabai，M．A．（1998）：Aminoacidcomposi−
genin rain and aerosolin the eastern Mediterraneantionofsoilorganicmatter．BiologyandFertilityofSoils，
atmosphere：Anassociationwithatmosphericdust．Jou−
26，235−242．
rnalof GeophysicalResearch，108（DlO），4320，doi：Short，S．KandHoldsworth，G．（1985）：Pollen，OXygenisotope
10．1029／2002JDOO2997．
COntentandseasonalitylnanicecorefromthePennyIce
McAndrews，］．H．（1984）：Pollen analysis of the1973ice core
Cap，Baf7inIsland．Arctic，38，214−218．
from DevonIsland Glacier，Canada．Quaternary Rer
Simon，M．and Azam，F．（1989）：Protein content and protein
SearCh，22，6876．
Synthesis rates of planktonic marine bacteria．Marine
McMurry，J．and Castellion，M．E．（1999）：Fundamentals of
Ecology Progress Series，51，201213．
OrganicandBiologicalChemistry．2nd Edition．Prentice
Takeuchi，NリKohshima，S．and Seko，K．（2001）：Structure，Hall，Inc．，NewJersey，544pp．
formation，and darkening process of albedo−reducing
Nakazawa，F．，Fujita，K．，Uetake，］りKohno，M．，Fujiki，T．，
material（cryoconite）onaHimalayanglacier：agranular
Arkhipov，S・M・，Kameda，T．，Suzuki，K．and Fujii，Y．
algalmat growing on theglacier．Arctic，Antarctic，and（2004）：Application of pollen analysis to dating ofice
AlpineResearch，33（2），115122．
COreSfromlowerlatitudeglaciers．JournalofGeophysi−
Wagenbach，D．（1989）：Environmentalrecordsin alpine gla−
CalResearch，109（F4），FO4001，doi：10．1029／2004JFOOO125．
Ciers．In The EnuironmentalRecordin Glaciers andIce
Olivier，S．，Schwikowski，M．，Brtitsch，S．，Eyrikh，S．，Gaggeler，
Sheets．（H．OeschgerandC．C．Langway，Jr．Ed．），pp．69q83，
H．W．，Ltlthi，M．，Papina，T．，Saurer，M．，Schotterer，U．，
John Wileyand Sons，New York，Tobler，LandVogel，E．（2003）：Glaciochemicalinvestiga−
Yoshimura，Y．，Kohshima，S．and Ohtani，S．（1997）：Commu−
tion ofanicecorefrom Belukhaglacier，Siberian Altai．
nityofsnow algaeon aHimalayan glacier．Arctic，Ant−
GeophysicalResearch Letters，30（19），2019，doi：10．1029／
arctic，andAlpineResearch，29，128L139．
2003GLO18290．
Yoshimura，Y．，Kohshima，S．，Takeuchi，Nり Seko，K．
Pollock，R．（1970）：What colors the mountain snow？Sierra
Fujita，K．（2000）：Himalayanice core dating with snow
Club Bulletin，55，18−20．
algae．JournalofGlaciology，46（153），335−340．
Reese，C．A．，Liu，KB．and Mountain，K．R．（2003）：Pollen disL