VariationsoftheNumberofEmbryosinMicrotusmontebelli
(MILNE−EDWARDS)*
YukibumiKANEKO BioIogicalLaboratory,FacultyofEducation,KagawaUniversity,Takamatsu,Japan
ABSTRACT.−Thedistributionofembryosinuterin¢horns,SeaSOnalchangesofbody Weightinpregnaptfbmales,andrelationbetweenthenumberofembryos,bodysize and age of pregnant fbmales,micro−habitat and season were describedin肋rotusmonlebelli.Number of specimens,Which were co11ected on ricefield,ricefieldin fallowandyoungplantation,WaS84.Therewasnostatisticallysignificantdeviation fromarandom distributjonofembryoswithintheuterus.Theaveragebodyweight
Ofpregnantfbmalessubtractingtheweightofembryoswasnotsignificantlydi鮎rent
duringayear.Theaveragenumberofembryosincreasedfromclass6to8andde− CreaSed fi.orn class9tollinage−grOupOfpregnantfヒmales.Therewasatendency
that theaverage bodyweight ofpregnantftmalesincreasedwiththeincreaseofthe
numberofembryosatthesameclassin age−grOup.Thenumberofembryosinthe SameWeightofpregnantftmalesonleveeswaslargerinSeptemberthaninMay.A
COmParison between two neighboring Octoberpopulations showed that the number
Ofembryosinthesamらbodyweightwaslargeronricefieldinfallowthaninyoung
plantation.Itissuggestedinthisvolethatthedifferences offoodqualitybrought aboutthevariationofthenumberofembryosofthesameageandbodysizeofpreg− nant fbmalesindi庁brentmicro−habitatsinthesameseasonand/orindi飴rentseasons in thesamemicro_habitat. IntroductionIthasbeenwe11knownthatthenumberofembryosorlittersizevaries和ithseason,
year,body size,POPulation density,▲habitat,altitude andlatitude.Severalpapers
havebeenpublishedinMicrotusn−Omwi1dpopulationsinEuropeandNorthAmerica
onth占variationofthenumber極relationtoage,SeaSOn,year,habitat,bodysizeand
latitude(JAMESON,1947;BEE正et∼al.,1957;BEER&MACLEOD,1961;KoTT&
RoBINSON,1963;NEGt)$&PINTER,1965;TAST,1966;EvANS,1973).IntheJapanese
丘eld vole,Microtus montebelli,however,ftw works have been carried out on the
Variationofthenumberofembryos,eXCePtthefbllowlngPaPerS:thenumberchanged
WithseasonduringayearinalocalityinJapan(MIYAOetdl.,1966;SHIRAISHI,1967;
ABE,1974;KANE竺0,1976)andwithhabitatsinneighboringOctoberpopulationsin
ayear(KANEKb,1978b).
ThepresentpurposeofthispaperistOdescribethedistributionofembryosin
uterinehorns,SeaSOnalchangesofbodyweightinpregnantftmales,andtherelations
betweenthenumberofembryos,bodysize−andageofpregnantfbmales,micro−habitat
andseasonintheJapanesefieldvoie,Microtusmontebelli(MILNE−EDWARDS).
*ContributionNo.40fromtheBiologlCalLaboratory,FacultyofEducation,KagawaUniversity.哺乳動物学雑誌 Vol.7,No.3
122
Materials and Metbods
Inthisstudy84specimens oftheJapanesefield volewere taken on rice丘eld,
ricefieldin fallow and young plantationin the fbllowlngfivelocalitiesinJapan:
Morioka,Iwate Pref.;Ry6zu,Niigata Pref.;Iwakura,Kyoto Pref.;Jinsekiand
Kurome,HiroshimaPref.;and Fukuoka,FukuokaPref.The specimens caught at
Iwakura,JinsekiandKuromewerethesameaspreviouslydescribedinKANEKO(1976,
1978a&b).Thespecimenswerecapturedbysnap−traPSWithpumpkinseedsasbait.
Al1fbmaleswereweightedandtheirbodysizesweremeasured.Thenumberofvisible
embryosinbothhornsoftheuteruswascountedwiththenakedeye,andthetotal
weightofembryoswasmeasuredincludingthefbetalmembranesanduterus.Body
weightsubtractingtheweightofembryoswas■uSedasthatofeachpregnantfbmale.
Theclassinage−grOuP Ofpregnantftmaleswasdeterminedbythemorphology
ofoccipitalprocessandridges,Whichwasdescribedinthepreviouspaper(KANEKO,
1978a).Sincetherewasagoodcorrelationbetweenbodyweightandthelengthofhead
andbody(KANEKO,1978a),bodyweightofpregnantftmaleswasusedasbodysize.
R(さSⅥ1ts and】Discussionl)Distributionofembryosinuterinehorns
Thenumberofvisibleembryosin40volesrangedn・OmtWOtOnineinthe1971−
72collectionsofIwakura,Kyoto.Theincidenceofunilateralpregnancywas7.5%
(3/40)and two specimens wereleft side pregnancies and one was right side.The
PrOPOrtionofbilateralpregnancieswithequalnumbersofembryosinbothhornswas
15.0%(6/40).Therewasnosignificantdeviationn・Omarandomdistribution(Poisson
distribution)ofembryoswithintheuterus,aCCOrdingtoax2testasshowninTablel
Tablel.Thedistribution ofembryosbetweenleft(Lt)andright(Rt)uterinehornsof 助川助∫椚♂〝Jg∂d〃atIwakum,Kyoto,197ト72. Difrbrenceinnos. No.ofvolesofembryosbetween Distribution betweenhorns Observed
(total) Expected no.(∽) borns Lt>Rt Rt>Lt (6) ′LU 5 1 7 0 1 1 1 2 0 つJ 4 2 0ノ 00 3 0 5 2 0 1 1 0.590 0.308 0.379 0.474 2.200 0.011 ズ2=3.962 O l つー3 4 5 00 5 4 0 1 7 ‘U つJ O O x,=difftrencebetweentheobservedandexpectednumberofanimals.Variance(sり=0.653・
(0.25<p<0.5).ThesamefindingwasobtainedinM.montebelliinFukuokaPre仁
bySHIRAISHI(1967)andinM.penn叩IvanicusandM.ochrogasterbyCoRTHUM(1967).
D Seasonalchangesofbodyweightinpregnantftmales
Inthe40specimensofIwakura,Kyoto,1971−72,thesma11estweightofpregnant
atIwakura,Kyoto,1971−72.
Month N Range(g) Meanj=SE(g)
May Sept. Nov. Jan. Mar. 32.5−38.9 29.9−41.5 27.7−38.5 22.6−29.2 26,0−37.1 7 4 7 3 ¢U l 3(5.13土.98 35.93土.83 32.26土1.50 25.27士2.00 33.04±1.25
ftmaleswas22.6ginJanuary,andthelargestwas4l.5ginSeptember(Table2).The
averagemonthlyweightoftheftmalesvariedftom25gto36g.Theaverageweight
WaS nearly equalthrough ayear exceptJanuary,When
Althoughtheaveragenumberofembryosdi鮎redstatisticallyamongmonthsexcept
between March and May(KANEKO,1976),the averageweightwas not significantly
di飴rent among six months.The mean bodylength ofpregnant fbmales did not
Change seasona11ythroughthebreedingseasonin ClethrionomysandbrsoniandApo−
ゐ∽〟∫α曙e〃fe〃∫(MIYAO,1969).
3)Thenumberofembryosagainstclassesinageqgroupofpregnantfemales
Thevisibleembryosappearedatclass6inage−grOuPintheIwakurapopulation,
Kyoto,1971−72.The average number ofembryosincreased ftom class6to8and
decreasedftomclass9tollduringayear(Table3),thoughthedifftrencebetween
Table3.Thenumber ofembryos under various classesin age−grOupin Microtus montebelliatIwakura,Kyoto,1971−72.The statisticaldifference between Class8andclass6was O.2<p<0.3and the difftrence between other classes exceptclassllwasp>0.6. No.indi鮎rentnumberclassesofembryos n p .1 u eS rO SS瑠 a e l Cag No.ofembryos (Mean±SE) 2 3 4 5 6 1 1 2 7 1 1 4 8 9 Total 12 4.6土.4 2 9 5.6士.8 1 4 5.8士1.2 6 5.2±.5 1 4 ‘U 7 00 q/ O 1 1 1 2 1 2 つん 2 2 1
Classesexceptclassllwasstatisticallyinsignificant・NEGUS&PINTER(1965)demon−
StratedinM・mOntanuSthatthelittersizeincreasedgraduallythroughthefifthcon−
SeCutivelitteranddecreasedthefbllowlnglitter.
Acomparisonofthenumberofembryos‘atthesameclassinage−grOuPShowed
that thelarge number occurredin September;While the smallone was fbundin
January(Fig・1)・Therefbre,thenumberofembryosatthesameclassinage−grOuP
Ofpregnantftmaleschangedwithseasons.
4)Relationshipbetweenthenumberofembryosandweightofpregnantftmales
InFig・2thenumberofembryosisplottedagalnSttheincreaseofbodyweight
哺乳動物学雑誌 Vol.7,No.3 124 。T 困刷冊SE脚 nU・ム 臼 ● ○
9 8 7 る 5 4 3 2 1
SOと芯≦山]○。OZ b ● ● 吐 血 血 nU O ・△ 03 4 5 る 7 8 9101112
ÅGE−GROUPFig.1.Scatter diagram of the relationship between classin age−grOup and the numberofembryosinMicrotusmontebelliatIwakura,Kyoto,197ト72. 0 9 8 7 d 5.4 3 2 1 SO≧垂烹〓〇.〇Z 0」AN,tr=−0.占d4) ム仙AR.=「=†0.る65】 ロ八AY け=†0.140) ●SEPT.け=†0.d35) O NOV.け=二0.089j ● ロ ●●臼 ●● 鎗 附● ●ム血● 00 ()A o dコ 0 ム 8 8 0 30 BODY WEJGHT(g) 40 Fig.2.Scatterdiagramoftherelationshipbetweenthenumberofembryosandbody WeightofpregnantfemalesinMicrotusmontebelliatIwakura,Kyoto,1971−72.
Bodyweight subtractingtheweight ofembryoswasusedasthat ofpregnant
females.The coe伍cient ofcorrelation(r)in each month was shownin this 負gure.
goodrelationbetweentheweightofmothersandthenumberofembryosina11the
SamPles during ayear(r=0.647).There was not so apparent correlation between the twoin each month(Fig.2),eXCePtin September(0.01<p<0.02).Heavier
ftmaleswithlargenumberofembryoswereobservedinSeptember,Whilelighterftmale
WithsmallnumberofembryosoccurredinJanuary.
bryos under various classesin age−grOupin MicrotLES montebelliatIwakura, Kyoto,1971−72. Mean*bodyweight(g)indifferentnumber Classesofembryos
Classes
111 age−grOuP Mean士SE 8 9 2 3 4 5 6 7 26.0 29.1 32.00土.97 40.3 35.01土1.85 39.1 34.98土2.23 35.72±.68 38.5 33.7 34.6 34.7 35.6 2夕.2 ′b 7 8 0ノ O 1 1 1 33.0 22.6 31.3 37.4 36.3 33.9 35.4 38.2 38.5 *NumberofspecimenswasgiveninTable3.thesame classin age−grOuP(Table4).Therewas atendencythatthe averageinq
creasedwiththeincrease ofthenumber ofembryos atthe sameclassinage−grOuP・
AccordingtoBEERetal.(1957),therewasasteadyincreaseinnumberofcorporawith
increaseillbodylenBth.Furthermore,they showed thatthe totalintrauterineloss
variedinverselywiththesizeofftmalesinM.pennsylvanicusandC.gqpperi.There−
fbre,the number ofembryos atthe same classin agergroupincreasedwith thein−
CreaSeOfbodysize.
5)Thenumberofembryosindi鮎rentmicro−habitats
The number ofembryos changed seasonallyinfourdifrerentmicro−habitats on
cultivatedland atIwakura,Kyoto,197ト72(Table5).The di飴rence of average
number ofembryos between di鮎rent micro−habitats was thelargestin September
amongfivemonths,thoughthedi鮎rencewasstatisticallyinsignificantin September
(0.1<p<0.2).In Septemberthe vegetation ofleveesinfallowlandswascomposed
mainly ofPob)gOnum thunbergii,Commelina communis and Cherus microiria,and
Gb)Cinemaxwasplanted on severallevees.Threepregnantftmalesweretakenon
fallowlandinJanuary(Table5),When fallowland was densely overgrown with
Withering Solidqgo altissimain abotlt2m height,and young Soli‘わgo altissima
Table5.Thenumber ofembryosin variousmicro−habitatsin difrbfent mOnthsin MicrotusmontebelliatIwakura,Kyoto,1971−72.Each尽gurerepresentsnumber Ofembryosofeachpregnantfemale≡collectedineachmicro−habitat.:Number inparenthesesshowstheaveragenumberofembryos.
Agricultul−al road‥‥D Fallowlandsdensely Grass−grOWnlevee Levee
inね1lowユand ユnrlCe ‥.B 丘eld….C 0VergrOWllWjth ふ痛血酢=油盆山那 ‥‥A 5,6(5.5) 4,5(4.5) 5 6,7,9,9(7.8) 5,5,5,5,5, 7 5,6,9,9(6,0) 4,4(4.0) 4,4,4(4.0) May 4,5,7(5.3) Sept. Nov. 2,4(3.0) Jan. 2,2,3(2.3) March 5 3,4,4,5,5, 5 5(4.3)
Vol.7,No.3 哺乳一動物学雑誌
126
sproutedatthebaseofthewitheredroots・
FromTable6,itislikelythattheaveragenumberofembryosindi能rentmicro−
habitats on ricefieldinfallowdidnot so greatlyvaryin May,June,August and
Octoberinsevera1localitiesinJapan.
A8C l ● l O ム︵ソ 8 7.〇 5.4 3
S2∝山王山L〇.〇Z
A ∞ム Ol ム○● ムム息ム血 130
40
BODY WE)GHT(g)
20
50
Fig.3・Scatterdiagramoftherelationshipbetweenthenumberofembryosar)dbodyweightofpregnantfemalesinMicrotusmontebelliindifferentmicro−habitatsat
Iwakura,Kyoto,1971−72.A;fallowiand densely overgrownwith Sb肋如 altissima,B:Grass−grOWnleveeinfallowland,C:Leveeinricefield・Open
symboIsrepresentspecimensinSeptember,andsolidonesshowspecimensin
May. 7 ′0 5 .4 3 つ‘ SO>出血王山 ﹄〇.〇Z 00 0 000 ■l 00● ○ ● ●● ● ● ● ● ●20 30 40 50
BODY WEtGHT(g) Fig.4.Scatterdiagramoftherelationshipbetweenthenumberofembryosandbody weightofpregnantfemalesin凡打crotusmontebelHinneighboringOctoberpopu− 1ations,HiroshimaPref.,1972.Opencirclesrepresentfemalesonricefieldinfallowandsolidonesshowfbmalesinyoungplantations.
コ ト.刊?寸 仇 M.刊﹁寸 p−ひ唱 Z 国S¶ロ召∑ 0−qd−亀ぎ JO∽払p佃銘 打︶ コ ︵∞.巴∽ゼゼ 寸.刊∽.れ ゼゼゼべれM m.刊○這 ぷp
鋸
Sき○竜∽OS月︶岩岳d点J月日nZ pdOh ︻空n︸ち0叫払く ︵巴 ︵?寸︶れhM や ︵﹁己れれれ れ音へ寸れMれM ︵?示︶ り.れれ寸 心小人ひJ rq︶ .sOh−q己ごOJOq己2=払巴璧ゐ﹄華 .︸眉q遥占JO叫∈竜dOひ︸8−ちじ書宍喜騙苫垂論ビd ざ.豊 トゼれれ ︵∞.寸︶ れーれ.れへれれれ寸 ︵竺寸︶り㌦.れれ寸寸 ︵∞.巴 トドぜ.れれ叫れれ p日月き0コd u叫0りAひ﹂ ︵U︶ く芭00琶 pロdlき○∃虎 口き○誌・S∽已0 ︵巴 ∽りS∽d︼叫 L≡き星g払 ﹂J診○とOS宕p p白虎lき○コd﹄ ︹く︶ やU−qd↑ M卜巴 さ羞=d琶nづ広 Nト仇︷ −月○︸00 3ひ∽月ヽ ︻トのー ︺sコ地コV コZ叉亘 Mトヘ︻ へ告nトムd∑ ぷ○叫旨∑ 重石○苫 き叫1日OJ哺乳動物学雑誌 Vol.7,No.3
128
Sincethenumberofembryosatthesameclassinage−grOuPlnCreaSedwiththe
increase ofbodyweight(Table4),COmParisons ofthe numberin the same weight
Weremadeamongdifftrentmicro−habitats(Fig.3).Thenumberinthesameweight
OnleveewaslargerinSeptemberthaninMay(B&CinFig.3).Itfb1lowsthatthe
number ofembryosin the same weight ofpregnant ftmales collectedin the same
micro−habitatchangedwithseasons.
Inthepreviouspaper(KANEKO,1978b),habitatdi飴rencesbetweenneighboring
Octoberpopulationsbroughtaboutthevariationoftheaveragenumberofembryos,
throughbothpopulationswereconsideredtobethesameinagecomposition・Figure
4showsthatthenumberofembryosinthesamebodyweightislargeronricefieldin
fallowthaninyoungplantation.
Summaringthesefindingsabovementioned,COmParisons ofthe same size and
age showedin Microtus montebellithat the number ofembryos changed primarily
Withthedi飴renttypesofmicro−habitatinthesamecollectingperiodandwithseasons
inthesamemicro−habitat.AlthoughEvANS(1973)showedinM.agrestisthatthere
WerenOSignificantdifrbrencesinpotentiallittersizesbetweenyoungplantationand
limestone grassland,NEGUS&PINTER(1966)revealedin M.montanus that small
dietary supplements ofeither sprouted wheat or acetone−ether extracts ofsprouted
Wheatelicitedimprovedreproductiveperfbrmance・Furthermor?,theydemonstrated
thattheimprovementresultedprimarilyinmorefiequentpostpartummaitings and
lower rates oflitterloss.Therefbre,itis suggested that the di鮎rences of food
qualitybroughtaboutthevariationofthenumberofembryosinthesameageand
Size ofpregnant ftmalesin diffbrent micro−habitatsin the same season and/orin
di飴rent seasonsin the samemicro−habitat.Acknowledgement
IwishtothanktoDr.Y.FuJIMAKIforreadingandcriticizlngthemanuscript・
摘 要 金子之史:ハタネズミにおける胎児数の変化 水田,水田休耕地および君命造林地で採集した84個体のハタネズミ妊娠雌について,胎児の左右 子宮内での分布,妊娠雌の体重の季節的変化,および胎児数と妊娠雌の体重・年令,捕獲された微小 棲息場所,捕獲時期との関係を調査した。 1.子宮内の胎児の分布のし方ほ,ポアソソ分布から統計的に有意な差を示さなかった。 2.胎児体重を差し引いた妊娠雌の体重は,1年間で有意な差は示さなかった。 3.平均胎児数は妊娠雌のage−grOup6から8にかけて増大し,9から10にかけて減少したが,統 計的に有意のある差ではなかった。 4.同一クラスのage一許Oupの妊娠雌で比較すると,胎児数の増加は妊娠雌の平均体重の増加に伴 なっていた。 5.畦で捕獲された同一体重の妊娠雌における胎児数は,5月よりも9月の方が大きかった。 6.同時期に採集した近接した2個体群内の同一体重の妊娠雌の胎児数ほ,水田休耕地の方が若令 造林地よりも大きかった。棲息場所の異なった採集時期,および同一時期に採集した異なった微小接点場所,によってもたらさ れる食物の質のちがいによって変化することが示唆された。
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