Miki-Hirosige, Hisako

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RIGHT:URL:CITATION:AUTHOR(S):ISSUE DATE:TITLE:A Study of Tropism of Pollen Tubesto the Pistils (VI) : Behaviour ofPollen Grains to Stigma of DifferentStages in DevelopmentMiki-Hirosige, HisakoMiki-Hirosige, Hisako. A Study of Tropism of Pollen Tubes to the Pistils (VI) : Behaviour ofPollen Grains to Stigma of Different Stages in Development. Memoirs of the College ofScience, University of Kyoto. Series B 1962, 29(1): 75-801962-05-26http://hdl.handle.net/2433/258657      MEMolRs ox" 'rnE CoLLEGE oF Sc!ENcE, UNIvERsn"y or- I<yorro, SEmEs B,                   Vol. XXIX, No. 1, Article 3 (Biology), l962       A Study of Tropism of Pollen Tubes to the Pistils              VI. Behaviour of Pollen G}'ains to Stigma                   of Di{{'eren'L S'Lages in l)evelopment                                  By                        Hisako MIKI-HIROSIGE             Botanical Institute, College of Science, University of Kyoto                          (Received July 28, 1961)   In the preceding papers it has been reported that some pistils in fiorescencestage contain certain active substances which induce the positive or negativetropism of pollen tubes and control the pollen germination and pollen tubegrowth CMiKi, 1954, 1955 ; MiKi-HiRosiGE, 1961a and b). It ls not clear, however,whether immature pistils contain these active substances or not. In Lilium,existence of these substances is confirmed more evidently in stigmas than inother parts of the pistiis. It is aimed, therefore, in the present study to seeeffects of tke active substances in different stages of stigma tissues in develop-ment on the tropism reaction of pollen tubes, pollen gerrnination and tke po}leRtgbe growth.   Stigmas of Li{itt•m iongiRorum in the following four stages, in respect tothe development of embryo sacs, were used.Stage A. Embryo sac mother cell stage.Stage B. Primary 2-4 Rucleate stage (meiosis stage).Stage C. Secondary four nucleate stage (florescence stage; early embryo sac         development).Stage D. Eight cell stage (four days after fiorescence; no pollinatioR).   Tropism reaction of pollen tubes, germination and tube growth of pollengrains spread around stigma slices in the four stages stated above were examiRed.Control tests were made without Åíhe stigma slices. Methods of preparing thestigma slices and of culturing polleR grains were described in details in theprevious paper (Mmi, 1954).   Result obtained in the present study is shown in the following table.   In this table, it is seen that while in Stage A the percentage of the positivetropism reaction of pollen tubes to the stigmas is nearly equal to that of thecontrol;in Stage B, the value is markedly higher than that in Stage A. Thepercentage reaches the maximum in Stages C and D. It must be emphasized76 Hisako MIi<I-HiRoslGETable 1. Tropism reaction, germination and tube growth of   pollen grains spread around the stigma slices in various   growth stages of L•il•iu?n longt'fior•u•nz.Growth stagesj"' -"""i COntrol e/o of germinated p'6'll'E'ff'{.. g.I.a.l.n..smm.....ma_. ... iii•ll'llll15111[i"ll.".IIIBI"...IIIi(ii.:11:ICr DTube length (mm)24 l 66o.3 l o.61L 6slL1...0.8O/e of positive tropism   Igo l    ]' "1'r"'ww""""..??..I mm?gL_l 5?wwl1.1'88840.786kere that in Stage A the styles and the ovaries do not affect the tropismreaction of polien tubes.   Both pollen germination and pol}en tube growth are promoted in StagesA, B,C and D in the presence of the stigma tissues (Table 1). The promo-tion reach the maximum in Stage C.   From the results stated above, it is concluded that, while the substanceswhich prornote the pollen germination and the pollen tube growth are containedin the stigma tissues in all stages in pistil development (Stages A-D), thesubstance whick induces the positive tropism appears in Stages B-D. It mustbe noted here that the substance which promotes the tube growth reaches themaximum in Stage C.   YAsuDA (1930) has reported that in the style of a fioral buds of Petuniathe rate of the tube growth of the pollen grains from the same fiower is rapidand ehe growth is normal in the bud stages as compared with those in thefiower-opening stage. He has expressed the view that the special substancewhich inhibits the growth of the pollen tubes is not yet pyoduced when thefiowers are still in buds. KAigzAKi (1930) and otkers have reported in Brassicathat, while the self iBcbmpatibility exists in fiower-opening stage, old fiowersof three or four days after tke fiorescence acquiye the force of self fertilization,and kave arrived at a coBclusion that the secretion of the special sttbstanceswill become wea}< as the physiological function is lost in the old fiower.tConelusion and Discussien   Results obtained in a series of the experiments carried out by the presentinvestigator show tkat the pistils of many ki.crher plants contain certaiR activesubstances which coBtro} the bel}aviour of the polien graiRs. If the inductionof positive tropism (MiKi, 1954, 1955), induction of negative tropism (Mii<i,1959; MiKi-HiRosiGE, 1961a), promotion of pollen germination (MiKi-HiRosiGE,1961b), inhibition of pollen germinatioB (MiKi-HiRosiGE, i961a and b), proinotioRof pollen tube growth (MiKi-HiRosiGE, 1961b) and iBhibition of pollen tubegrowth (MiKi-HiRosiGE, 1961a) are caused from different substances respectively,six substances may be assumed. These substances are summarized in Table 2.Table2. Substancescontained in pistiltlssueswhichaffect thebehaviours ofpollen grains.Characters /    ofIsubstances ,Plant names    of  pistilsi PIant names llreacted p8i(en gramsl         'lExtracted I         I solvent i         ir"'r--   He.t ' Diffusion lg,er,M.rf.b,iigt.Ytttttt fiieP:Etlww.1.. gFa;?sls.1/-r,ifiglE,yl2se' Induction of    posltlve    troplsm-' in'a'Glll6'n-6"l""'   negative    trQplsm-' "r6-O-tl6"' ''6k' 'nv     pollen...gg.{{I.}lnglip.llma•• Li• tiuinlongiflorsc}n: :Liliec?n longifiorum 'Camegtia sinensisunstableunstable.?.I9.VigstablediffusiblepermeableCameltia sine"sis..erylee.uia CaMelliaobconicaSi?ZettSiS*prim"la o.bcg7iKfZCameliia sinensis diffusible•..A.l'.ff_Y.?}IP.l.9...._ .. diffusibleg.iB'Gilg.'fe.''p5}'m'"556''i, ''tirl}?i.};.ieggPiel".III99F.M..9.B.?i9P9f.M.9aPi9....gis.tli• rtlgE.er3gi.gy,sc.wa.,.distil. water 1 ttt t.ttt.tt ttttt tt. I           idistil. water !           I.gl.9.t3]• NYat9r idistii. waterchloroformdistil. yatef. ,.ai5tii. yager ,l MIK.{, MIKI,1 M.Åí1<!-I{IROCSIC-. F-,  l961a .t MI!<I, MII<I-HIIIOSICrE,  l961 b'M'i'i(I.KIRoslC,tl,"'mm  1961 a Miig-Hu?osicirt,  l961 b Mll<I.HIRoSIGE,  1961 bLiterature  1954  1955  1959'Litium lo}tgifierecinI[ iPPeastrunz  itybridunt' i'll:-Ia2'k'' obconica'ec:.be'l,ss..#,.'j 1,;,,agzef3Es[-)1'l1.lPromotion of i Liiitt"t tO}igifiereciiipo,i;s?.,`,"be lthb,glbg'f:AiÅít;;f'""pmwwInhibition ofe9U'?g,,lec.er"...er.iilzff.i.a.um.gbfe.nica Lilium tongifioncnt /"'ll. '1'l 'i'b'b's' tr.,,i ' ''''t't't''t,'- ItybridumPrimzda obconicai : E iLilium longiflorumHiPPeaStrUM   hYbridUM.Ang.e"(c(iglzl.ggijz.maju.s..Ligiu7n longiflorui}t stabie stable stable stablel stable.fteuaf,lg.,",:iww."...., primuta obconica lstablestablediffusib}ediffusiblediffusible' a,'itl'gl,6"i5"'1-g-if, ilglg---permeabie impermeaPle imperrneable..?9fill.P.91.!9........ permeableimpermeable           MJ.I<I-HIROSXGE,distil. Water i 1961 a\' Steamedpistil.> m 8 EX g B 8 Y. ca p g oru o"-" " E st cr o ca ti o el tr o w rn' pt/' NNca < M li78 Hisako M!iq-HIRoSIGE   In spite of the above assumption, it is not established definitely in thisseries of experiment whether all of these substances are differen.t or not. Someconslderations oR this question are given as follows :   '1) TroPism indu.ci}?.g s"bsta•nces. Two substances which induce the tropismreaction of pollen tubes are found in the pistil tissues ; one induces the positivetropism and the other does the negative. Theformer substance is found in thefresh pistils of many phanerogams such as Ltli'es}}t, HePPiast?'um and Narcisstts(Mii<i, 1954, 1961), and the latter is found in the fresh styles of Pri7nula obconica(MiKi-HiRosiGE, 1961). In Cameglia, however, the fresh pistil tissues inducethe positive tropism, while the tissues of pistils treated at 990C induce thenegative tropism (MiKi, 1955). Therefore, it is concluded that the substancewhich induces the positive tropism in Camellia and Lilium is metastabJe toheat. Moreover, this substance diffuses through both the collodion and ce}lulosemembranes. Contrary to the above substance, the substance which inducestke negative tropism is stable to heat and does not diffuse through tkesemembranes.   Considering the facts stated above, it is quite safe to conclude that thesetwo substances are quite different. In the pistils of Ligium, it is assumed tkatonly the substance wkich induces the positive tropism in coBtained (Mu<i, 1954),while the style of Primula obconica contains only the negative tropism inducingsubstance. In Camellia sinensis, however, lt is assumed that both the positiveand the negative tropism inducing substances are contaiBed is, the pistils, andthat the former is more active in fresk tissues. When the pistils are heated,only the former substance is destroyed while tke Iatter exhibits its activity(MiKi, 1955).   2) Germi'nation and tttbe growth controlli•ng szcbsta•nces. In many plants suchas Lilit{m, Narcissus, Camellia and HePPiastrum, both ihe germination and thetube growth of pollen grains near the pistils of the same species are promoted,but this promoting effect is not recognizable in the pollefi grains fiot close tothe pistils (MiKi, 1954, l961).   Then, it is questioned whether tke tropism controlling substances aredifferent from the germination and the tube growth controlling substances ornot. The results of the experiment reported iR this paper suggest a strongprobabllity that these two substances are different, because the substances whichpromote the germination and tube growth are contained in stigma in all stagesin the course of pistil development, while the substance whick induces thepositive tropism appears on}y in the later stages of the pistil development.   Then comes the question that wltether the germination contro}ling sub-stance differs from the tube growth controlling substance or not. Unfortunately,this question is not clearly soived in this series of experimeRts, but the resultobtained in the experiment reported iR 1961 probably supports a view tkat thegerrniRation and the tube growtl are controlled by differeRt active substances               A Study of Tropism of Pollen Tubes to the Pistils, VI 79(Mii<i-HiRosiGE, 1961b). In this experiment it is confirmed that, when thepol}en grains of HiPPeast3'ynt are spread around the style siices of CSivia, thepollen germination is ma.rkedly arrested while the tube growth is slight}ypromoted. Moreover, when the pol}en graims of Hi ppeast "i ttm are spread aroundstigma slices of L. ;oGtgiflorum, the tube growth is arrested while the germina-tion coethcient is almost equai to that of the control.   There is aBother questioR whether the poilen germination promoting sub-stance and the pollen germlnation inhibiting substance are same or not. Theresults of the experiment carried out by the present investigator oR Primtttaobconica are regarded suthcient to give a certain clue to solve this question(MiKi-HiRosiGE, 196ia). It is observed in this plaRt that in the pollen grainsclose to the ,style slices the germination of pollen grains are inhibited, whilein the pollen grains away from the style slices the germination is promoted.It is fair, therefore, to assume that both the lnhibition and the promotion arecaused from different concentrations of an active substance. At least in Primula,however, there is no reason to infer that the germination promoting substanceand the germination inkibiting substance are different. While in Primula thegermination controlling substance does not diffuse through the cellulose andcollodion membranes, in Lilium and Htppeastrum the germination promotingsubstance diffuses through these membranes. Tkerefore, it is assumed that tkegeymination promoting substance in Lilium and H•ippeastru•m and tke germinationcontrolling substance in Pr•iinttta are differeBt. Then, it is inferable that thesubstances wkich promete the germination are not always the same.   Similar relation between the tube growth controlling substance in Primulaand the tube growth promoting substance in Lilium and. Hippease?'um is observed.Therefore, it is assumed that the tube growth promoting substaRce in Liliumand MPPeastrum and the tube growth controlling substance in Primula aredifferent.   In the series of this experiment, it is not observed tkat the pollen grainsor the pollen tubes show a special behaviour around a special part of thepistil. For example, in Lilium the tropism reaction around stigma slices is$tronger than that around the style or the ovary slices, while in Camellia sinensisit is stronger around style slices than that around the other.   Genera}ly speakiRg, it is concluded that the positive tropism reaction, thepromotion of pollen germinatioB and that of the poilen tube growth are seenwhen the relationship between the plaRt from which the pistil is taken andthe plant from which the pollen grains are taken is near, but this teRdency ishardly observed when the relationship between the two plants is remote. Itmust be Roted here, ltowever, that a few exceptional cases are observed ikthis series of the experiment. These facts may be explained as follows : Someactive substaRces, which control the tropism reaction, the germiRation and thetube growth, are contained together in the pistils of higher plants, and thebehaviours of the po}len grains to the pistils are caused from the difference80 . Hisako }vlH<I-HiRoslGEin localization of the active substances, difference in the time in whiek theactive substances are produced, difference in concentrati:on of these substancesand other factors.                               Summary   1. Tropism reaction, germination and tube growth of pol}en grains spreadaround the stigma slices are studied in different stages in pistil developmentof Lilium go7i.gifioru]it.   2. Substances that promote the pollen germination aRd the pollen tubegyowth are detected in the stigma tissues in all stages of development, whilethe substance whiclt induce th.e positive tropism is found only in the later   3. Relation among the substances which control the tropism, the germinatioRand the tube growth of pollen grains is discussed.   The author takes pleasure in expressing ker sincere appreciation for help-fttl suggestions throughout this work te Professor N. SmNKE and Dr. K. KAT6of Kyoto University.                           Literature CitedJosT, L., 1907. Bot. Ztg., 65.KAi<IzAKI, Y., 1931. Jap. Jour. Bot., 5.Mii<r, H., 1954. Bot. Mag. Tokyo, 67.mm......t.ttttttt-ttt-w  1955. Ibid•, 68•---ttttt----t-t-  1959. Mem. Coll. Sci. Univ. Kyoto, (B), 26•-'ntt''t''r  1961. Ibid•, 28•Mii<i-HIRosiGE, H., 1961a. Ibid., 29.----------  1961b. Ibid., 29.YAsuDA, S., 1930. Bot. Mag. Tol<yo, 44.

A Study of Tropism of Pollen Tubes to the Pistils (VI) : Behaviour of Pollen Grains to Stigma of Different Stages in Development

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A Study of Tropism of Pollen Tubes to the Pistils (VI) : Behaviour of Pollen Grains to Stigma of Different Stages in Development

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