Kato, Kazuo

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RIGHT:URL:CITATION:AUTHOR(S):ISSUE DATE:TITLE:Chromosome Behaviour in theInterkinesis (II) : Observation ofArchesporial and Spore MotherCells in Psilotum nudumKato, KazuoKato, Kazuo. Chromosome Behaviour in the Interkinesis (II) : Observation of Archesporialand Spore Mother Cells in Psilotum nudum. Memoirs of the College of Science, Universityof Kyoto. Series B 1963, 30(3): 53-571963-12-20http://hdl.handle.net/2433/258687MEMolRs OFTHE COLLEGE Vol. XXX. oF SCIENCE, UNIVERSITY OF KYOTO,No. 3, Article 5 (Biology), l963SERIEs B,Chromosome Behaviour in the InterkiResis     II. Observation of Archesporial and Spore         Mother Cells in PsiZotum nudumBotanical              By         Kazuo KAToInstitute, Faculty of Science,  (Received September 23, Kyoto1963)University    Since the first paper of the series on the observation of the iRterkinesis in'Tradescantia (KATo, 1935) being published, years have passed without publishingof thls series, owing to the Second World War and uncler circumstances of theauthor. Observations made at that time in other plants wil} be reported iR succeediRgpapers.    In the present investigation the nuclear structure in the interkinesis in PsilotMmwas observed in connection with the hericali) structure of chromosomes. Somecomparison of the interkinetic nucleus was also made with the nuclei iR the inter-phases (last pre-meiotic and post-meiotic divisions) and with the periplasmodiumnucleus.                           Material and Method    As materials, archesporial and spore mother cells of Psiiotum nudum wereused. The cells wei:e stained with the ordinary aceto-carmine method. In somecases SAx's ammonia-alcohol mixture (SAx and SAx, 1935) was a}so applied beforestalning with the aceÅío-carmlne. In the observation of living state liquid paradin andsaccharose solution were used as media.                                Observation    Last Pre-meiotic metaPi?ase to teloPg?•ase. Archesporial cells in Psilotum areundergone a few pre-meiotic divisions before they enter into the meiotic division, andthe cells originated from an archesporial cell are in groups. Of these grouped cellsthe maximum number was observed to be 8.   1) In place of spirals which have often been misused, the term helices are used inthe present paper. 54 Kazuo KATo    ln the last pre-meiotic metaphase the chromosomes are slender as in ordinarysomatic divisions. In Fig. 1, in a chromatid (half chromosome) chromonema heliceswith small somatic coils are seen in the matrical substance. In most chromatidsthe chromonemata appear in two helices lying side by side along Åíhe chromatldlength and iR others they appear as a single helix.    Fig. 2 shows a telophase nuc]eus, where the chromosomes take polar orientationleaving a clear space at the pole of the nucleus (pole field). In this figure thechromosomes are revealed as solid threads otherwise than internal strttcture. Inthese solid threads the two chromonema helices observed in the metaphase areconcealed, so closely associated together to appear as a single helix. This chro-monematal association might be caused by the Iengthening out of chromosomes iB iatetelophase, as pointed out by ATwooD (1937) in the iast pre-meiotlc telophase ofGaillardia aristata.    Last Pre-meiotic interPhase. In the interphasic nucleus in the last pre-meioticdivision, the thick solid hellces observed in the preceding telophase are transformedinto the form of herical chromonemata with small coils (somatic type). With thesechromonema helices the nucleus is densely fi}led, the polar space aRd spaces betweenthe chromosomes are lost. The chromonema helices are }oosely relaxed aRd in-tricately expand in the nucleus,:so that it is not possible to distinguish Åíhe chromonemaherices which have beeR derived from a single chromosome (Figs. 3 and 4).    The diffuse iRterphasic nucleus in Psilotum presents a striking contrast to that inthe same stage in Bellevalia ronzana (OEm.KERs aRd EBERLE, 1957), iR which thechromosomes are less unravelled and the nuclear helices with different diametersalong their length are disposed sparsely in the nuc!eus.    Fivst meiotic metaPl?.ase to tetoPhase. In P. nudum, about 50i) thick and shortchromosomes of somewhat different size are found in the first metaphase. IR maRyorganisrns, especially in plants, compact and condenced chromosomes in the firstmetaphase are observed being composed of two distinct helices, major and minor;the minor helix is coiled again into the major helix with larger coils (KuwADA, 1932,1935, 1939; KuwADA and NAKAMuRA, 1933; OuRA, 1936; GEiTLER, 1938; MATsuuRA,1938; CoLEMAN and }IiLLARy, 1941; DE RoBERTis et al., 1960).2) In the metaphasechromosomes in this plant the chromonema helices are coiled into helices of themajor type, in which the minor helices are usually concealed (Fig. 5). The majorhellces can be recognized already in diakinesis, where a conspicuous shortening ofchromosomes is taken place. The major hellces can be relaxed, when ammonia-alcoholmixture ls applied as pretreatrr}ent before staining with the aceto-carmine. IR Fig. 6,the out-line of the chromosomes (chromosome matrix) ls lost, and coiling of themajor Åíype is evident as continuous helices. In liquid paraffin, the chromosomesappear as homogeneous bodies, showing no interRal structure. The same homogeneousappearance of the chromosomes is presented in a 15.(b/ saccharose soiution. IB this    1) According to OKABE (1929), n-52.    2) For further literature on the double-coiled structure of chromosomes refer toMANToN (1950) and RucH (1950).                    Chromosome Behaviour in the Interkinesis, II 55case, however, the spore mother cel}s show a slight plasmolysis in a meantime.    The massive form of the chromosomes is maintained from the metaphase untilthe daughter nuclei are formed in the telophase. When the nuclear membrane isformed, the major helices begin to unravel and the shape of individual chromosomesbecome less evident (Fig. 7). With these changes one nucleolus makes lts appearance.At early telophase the formation of a separating plate or wall of tb_e cel} is firstvisible in liquicl paraffin. However, this plate formation is always non-visible inaceto-cat'mine preparations.i)    Interkinesis. In the interkinesis the major helices continue a further unravellingand the minor helices concealed in the major ones during previous stages becomeevident. A transitional stage from the major coiling into the minor one is shown inFlg. 8. In this nucleus not only the minor helices but also the major ones are seen.In Figs. 9 and 10 the major helices are not completely unravelled and they arerecognizable as such here and there in the nucleus. In Fig. 10, in the lower part inthe fiucleus, it may be seen that two minor helices of a chromosome, to be separatedin Åíhe second division, come loose from each other except for the attachment point ofspindle fiber. The same feature has been observed in the same stage of Tradescantiareflexa (KATo, 1935). In Fig. 11, the unravelling of chromosomes is reached iÅísmaximum. I!i this nucleus the chromonema helices with minor coils are mostevident. Although clear spaces are seen between 'territories which have beeR occupiedby chromosomes, it is dithcult to discriminate chromonema helices derived from asingle chromosome.    In comparing with the interphaslc nuclei in the pre- and post-meiotic (see b}ow)divisions, the chromonema helices in the interkinetic nuc}eus are coarsely distributed.In Psilotttm a general appearance of the interkinetic nucleus, in iÅís maximurnchromosome unravelling, resembles to that in Tradescantia veRexa (KuwADA, 1932;KATo, 1936). However, with regard of presence of the relic major coils, theinterkinetic nucleus in Psilotum might be classified as to be a member of Liliscmtype (SiNKE, 1934; KATo and IwATA, 1935).    As the stage advances toward the end of the interkinesis, the nucleus siÅíuatedclosely to the cell periphery at the poles of the mother cell, moves toward the centralin the daughter cells.    Fig. 12 shows an early prophase in the second division, in which the chromo-nemata begin to thicken and the minor coils become iess evideRt, showing more orless larger coiling of prophasic Bttclear threads like in the spiral stage ln somaticmitoses (cf. KuwADA and NAKAMuRA, 1934).    Second metaphase. The chromosomes in this stage are somewhat slender shapeas compared with those in the first metaphase, and they appear to be composed ofhelices of the minor type (Fig..13). In llquid parathn the chromosomes appear tobe homogeneous as in Åíhe first metaphase.    Post-meiotic tnterpl?ase. In the post-meiotic interphase which follows the second   i) 'in th-e Ia'st prellll'eio'tiE' aivislo-E','the separating waii can be seen in aceto-carminepreparations.56 Kazuo KATodivisioR, the nucleus is filled with the minor he}ices (Fig. 14). These minor helicesare similar iR coiling to those found in the last pre-meiotic interphase.    PeTiPlasmodium nucleus. A tapetal periplasmodium nucleus, nature of whichis nutrkive, is shown in Fig. 15. In this nucleus the chromosomes are in the helicalform with small somatic coils. Among these helices pycnotic bodies are observed.Such pycnotic bodies can not be observed in the interphasic nucleus in the last pre-meiotic division.                                 Summary    1) In Psiiotttm mtdztm, the nuclear structure in the interklnesis was observed,taking its structural connecÅíion to other stages into consideration.   2) IR the interkinetic nucleus, the meiotic chromosomes composed of major andminor helices are transformed into their constituent helical chromonemata. At itsculminating stage the nucleus is fi}led mostly with the minor helices, with somerelics of the rnajor helices.    3) In comparing the Ruclear structure in the interkinesis with that ln thesomatic interphases (pre- and post-meiotic divisions), texture or pattern of the chromo-nema helices ls coarser in the former than in the latter. In the periplasrfiodium nucleuspycRotic bodies are observed among the chromonema helices.Literature CitedATwooD, S., 1937. La Cellule, 46: l89-410.CoLEMAN, L. C. and B. B. ffiLLARy, 1941. Amer. Journ. Bot., 28: 46(l--ti69.DE RoBERTis, E. D. P., W. W. NoviNsKi and F. A. SAEz, 1960. General Cytology           Philadelphia and London.GEiTLER, L., 1938. Chromosomenbau. Protoplasma-Monographien, 14. Berlin.KATo. K., l935. Mem. Coll. Sci. Kyoto Imp. Univ., Ser. B, 10: 251--262.       and J. IwATA, 1935. Ibid., 10: 265--273.KuwADA, Y., 1932. Bot. Mag., Tol<yo, 46: 307-310.t._ ,i935. Cytologia, 6: 308--313._. _,1939. Cytologia, 10: 213-256.     .nd and T. NAi<AMuRA, 1933. Mem. Coll. Sci., Kyoto Imp. Univ., Ser. B, 9:MANToN, I., 1950. Biol. Rev., 25: 486-5e8.MATsuuRA, H., 1938. Cytologia, 9: 2t13-248.OEHLKERs, F. and P. EBERLE, 1957. Cytologia, 8: 351--363.OKABE, S., 1929. Sci. Rep. T6ho{<u Imp. Univ., Fourth Ser., 4: 373--379.OuRA, G. 1936. Zeitschr. wiss. Mikros. [Irech., 53: 36--37.RucH, FR., 1950. Chromesoma, 3: 358--392.SAx, H. J. and K. SAx, 1935. Journ. Arbor., 16: 423-439.SiNKE. N. 1934. Mem. Coll. Sci., Kyeto Imp. Univ., Ser. B, 9: 367--392.(third ed.)129-139.Chromosome Behaviour in the Interkinesis. II 57                             Explanation of Plates                                    Plate I   Figs. I--l5 are photomicrographs taken from aceto-carmine preparations, using ZEiss'sapoch. imm. 2mm. and comp. oc. 12.   Fig. 1. Chromosoines in the last pre-meiotic metaphase. .   Fig. 2. Last pre-meiotic telophase.   Fig. 3. Interphase nucleus in side view.   Fig. 4. The same in polar view.   Fig. 5. I-metaphase chromosomes, showing inajor helices.   Fig. 6. Ernravelling of I-metaphase chromosomes pretreated with SAx's inixture,showing major he]ices.part                                Plate IIFig. 7. I-telophase, beginning of unravelling of major helices into minor ones.Figs 8 and 9. Interl<inesis. Major helices are more evident than minor ones.Fig. 10. The same. Two miner helices of half chromosomes are seen in the lower of the nucleus.Fig. 11. Mid-interkinesis, major helices being unravelled to the maximum point.Fig. 12. Early II-prophase, slaowing thickening of nuclear helices.Fig. 13. Chromosomes in II-metaphase.Fig. 14. Interphase in post-meiotic division.Fig. I5. A periplasmodium nucleus, showing diffuse structure of metabolic stage.Meni. Coll. Sci., Univ. Kyoto, Ser. B, Vol. XXX, No. 3, Art. 5, Pl. I.tl#rv••.;JrwA, {1: ltLNrw .,X "es " +t3..g,Lfu' feq;'2NX;' -'t'-' ".N"'srl•'s--[. l•Ii "'"t.S;.'{gX4''tl'ij -tLS'...- 't'-za,   -.:.k{[tt.. ...l..-i..    K. KATOMem. Coll. Sci.. Univ. Kyoto. Ser. B. Vol. XXX. No. 3, Art. 5. Pl. II't nP7''  i' ''   -' # '-t { N.., :N" .. '-. .''-'1 "t 't e '' '' - "r''''  -,'  Il1--':ees""vi' "t'.r..'x' l .wS   " " -" -"  ? ' .s -,."i t .  --' /     ' - -' -L:L-r :)s•1';.,'IS,F li.4 s,-,'rwh'pu,'t' t-- ' bts-  l•-, 9'f. E r .:rsr'rS1 i:...r1.. ".'1• `i.iilt. ko }'N a ,•f V" !. .1tt-Å}S.t s9..x'-.' xtg1. 1..V 1+22 fS.di-) sk3    E" cg IJ"'l5-K. KATO

Chromosome Behaviour in the Interkinesis (II) : Observation of Archesporial and Spore Mother Cells in Psilotum nudum

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Chromosome Behaviour in the Interkinesis (II) : Observation of Archesporial and Spore Mother Cells in Psilotum nudum

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