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E.Olmo C. A. Redi

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ChromosomesToday

Ettore Olmoearlo Alberto RediEditors

Springer Basel AG

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Edited by

Prof. Dr. Ettore Dlmo Istituto di Biologia e Genetica Via Brecce Bianche 60131 Ancona Italy

Prof. Carlo Alberto Redi University of Pavia Dipartimento di Biologia animale / Centro di Studio per l'lstochimica - CNR Piazza Botta 9 27100 Pavia Italy

A CIP catalogue record for this book is available from the Library of Congress, Washington D.c., USA

Deutsche Bibliothek Cataloging-in-Publication Data Chromosomes today : volume 13/ Ettore Dlmo ; Carlo Alberto Redi, ed .. - Basel ; Boston; Berlin: Birkhauser, 2000 ISBN 3-7643-5799-1

The use of registered names, trademarks etc. in this publication, even if not identified as such, does not imply that they are exempt from the relevant protective laws and regulations or free for general use. ISBN 978-3-0348-9587-3 ISBN 978-3-0348-8484-6 (eBook) DOI 10.1007/978-3-0348-8484-6

This work is subject to copyright. AII rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, re-use of illustrations, recitation, broadcasting, reproduction on microfilms or in other ways, and storage in data banks. For any kind of use permission of the copyright owner must be obtained.

© For the book title ("Chromosomes Today, Volume 13") and the series title ("Chromosomes Today Series"): The Drganizing Committee of the 13th International Chromosome Conference, Numana, Ancona, Italy 1998 © 2000 Springer Basel AG Originally published by Birkhăuser Verlag, P.D. Box 133, CH-4001 Basel, Switzerland in 2000 Softcover reprint of the hardcover 1st edition 2000 Printed on acid-free paper produced from chlorine-free pulp. TFC Cover design: groflin. Graphic Design, Base!. www.groeflin.ch

ISBN 3-7643-5799-1

987654321

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Contents

List of contributors IX

Dedication XXI

Preface XXIII

Introduction

Chromosomes yesterday: a century of chromosome studiesErnesto Capanna 3

Part one: Chromosome organisation

The prometaphase bands of human chromosomes: compositionalfeatures and gene distributionGiorgio Bernardi, Concetta Federico and Salvatore Saccone 25

The functional and structural organization of DrosophilaheterochromatinSergio Pimpinelli 29

RNA, genes, genomes and chromosomes: repetitive DNA sequencesin plantsI.S. (Pat) Heslop-Harrison .... . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 45

Color plates I-VII

Non-B DNA structures spatially and sequence-specifically associatedwith individual centromeres in the human interphase nucleusMizuki Ohno, Toyoaki Tenzen, Yoshihisa Watanabe,Tetsushi Yamagata, Shigehiko Kanaya and Toshimichi Ikemura 57

Part two: Chromosome function and behaviour

Chromosomes in mitosis: chance and checkpointR. Bruce Nicklas 73

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VI Contents

Genetic control of B chromosome transmission in maize and ryeMaria J. Puertas, Guillermo Jimenez, Silvia Manzanero,Mauricio A. Chiavarino, Marcela Rosato, Carlos A. Naranjo andLidia Poggio 79

Igf2 imprinting in development and diseaseWolf Reik, M. Constancia, W Dean, K. Davies, L. Bowden,A. Murrell, R. Feil, J. Walter and G. Kelsey 93

Part three: Meiosis

Seeing meiotic chromosomes without seeing themNancy Kleckner 107

Meiosis in budding yeast and in multicellular eukaryotes - similaritiesand differencesJosef Laidl 123

Meiotic pairing sites and genes involved in segregation of the X and Ychromosomes of Drosophila melanogasterBruce D. McKee, Chia-sin Hong and Siuk foo 139

Part four: Chromosome evolution

Genome evolution in TriticeaeBelen Maestra and Tomas Naranjo

Karyotypic evolution in Neotropical freshwater fishLurdes F de Almeida-Toledo, Fausto Foresti andSilvio de A. Toledo-Filho .

Repetitive DNA as a tool to study the phylogeny of cold-bloodedvertebratesTeresa Capriglione

Comparative gene mapping permits new insights into karyotypeevolution in mammalsGunther Rettenberger, Walther Vogel and Horst Hameister .

Color plates VIII-X

Part five: Medical and cancer cytogenetics

155

169

183

195

Chromosome and gene alterations in human cancers in relation to agingBernard H. Dutrillaux 207

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Contents VII

Xp contiguous gene syndromes: from clinical observation to diseasegene identificationBrunella Franco and Andrea Ballabio 225

Genetic and cytogenetic studies in inherited cancer: Li-FraumenisyndromeJennifer M. Varley, John M. Boyle, Louise A. James, Gail McGown,Mary Thorncroft, Martin Greaves, Anna M. Kelsey, D. Gareth R.Evans and Jillian M. Birch 245

Color plates XI-XVIII

Part six: Mapping and new methods

Comparative chromosome paintingMalcolm A. Ferguson-Smith, Patricia CM. O'Brien, Willem Rensand Fengtang Yang 259

High resolution FISH reveals the molecular and chromosomalorganisation of repetitive sequences of individual tomato chromosomesJ. Hans de Jong, x.-B. Zhong, P.P Fransz, J. Wennekes-van Eden,£. Jacobsen and P. Zabel 267

Species relationships, introgression and physical mapping in Loliumperenne x Festuca pratensis hybridsIan P. King, WG. Morgan, 1.P. Armstead, J. King, T.A.H. Jones,J. V. Nash, R.N. Jones, J.A. Harper, L.A. Roberts, Steve Taylor andH.M. Thomas 277

Smallness: gain and loss in plant chromosome researchKiichi Fukui, Nobuko Ohmido and Toshiyuki Wako 287

Conclusion

The future of chromosomesHerbert C Macgregor 305

Subject Index

Synonymous words

315

320

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IX

List of contributors

Almeida-Toledo, Lurdes F. deDepartamento de Biologia, Instituto de Biociencias, Universidade de SaoPaulo, Caixa Postal 11461, 05422-970 Sao Paulo, Brasil, Tel: +55 11 8187584, Fax: +55 11 8187553, E-Mail: [email protected]

Armstead, I. P.Institute of Grassland and Environmental Research, Plas Gogerddan,Aberystwyth, Ceredigion, SY23 3EB, Wales, UK

A. Toledo-Filho, Silvio deDepartamento de Biologia, Instituto de Biociencias, Universidade de SaoPaulo, Caixa Postal 11461,05422-970 Sao Paulo, Brasil, Tel: +55 11 8187584, Fax: +55 11 8187553, E-Mail: [email protected]

Ballabio, AndreaTelethon Institute of Genetics and Medicine, San Raffaele BiomedicalScience Park, Via Olgettina 58, 1-20132 Milano, Italy, Tel: +39 02 21560202, Fax: +390221560220

Bernardi, GiorgioLaboratorio di Evoluzione Molecolare, Stazione Zoologica "Anton Dohrn",Villa Comunale, 80121 Napoli, Italy, Tel: +39-81-5833300, Fax: +39 812457294, E-Mail: [email protected]

Birch, Jillian M.CRC Paediatric and Familial Research Group, Royal Manchester Children'sHospital, Pendlebury, Manchester M27 1HA, UK

Bowden, L.Laboratory of Developmental Genetics and Imprinting, DevelopmentalGenetics Programme, The Brabraham Institute, Cambridge CB2 4AT, UK,Tel. +44 1223496000, Fax: +44 1223496030

Boyle, John M.CRC Section of Molecular Genetics, Paterson Institute for CancerResearch, Wilmslow Road, Manchester M20 4BX, UK, Tel: +44 161 4463062, Fax: +44 161 4463062

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x List of Contributors

Capanna, ErnestoDipartimento di Biologia Animale e dell'Uomo, Universita di Roma "LaSapienza", Via A. Borelli 50, 1-00161 Roma, Italy, Tel: +3964991 8008,Fax: +3964457516, E-Mail: capanna@ axrma.uniromal.it

Capriglione, TeresaDipartimento di Biologia Evolutiva e Comparata, Via Mezzocannone 8,1-80134 Napoli, Italy. Tel: +39 081 2528946, Fax: +39 081 2528913,E-Mail: [email protected]

Chiavarino, Mauricio A.Departamento de Biologfa, Facultad de Ciencias Exactas y Naturales,Universidad de Buenos Aires, 1428 Buenos Aires, Argentina

Constancia, W.Laboratory of Developmental Genetics and Imprinting, DevelopmentalGenetics Programme, The Brabraham Institute, Cambridge CB2 4AT, UK,Tel. +44 1223,496000, Fax: +44 1223496030

Davies, K.Laboratory of Developmental Genetics and Imprinting, DevelopmentalGenetics Programme, The Brabraham Institute, Cambridge CB2 4AT, UK,Tel. +44 1223,496000, Fax: +44 1223496030

Dean, W.Laboratory of Developmental Genetics and Imprinting, DevelopmentalGenetics Programme, The Brabraham Institute, Cambridge CB2 4AT, UK,Tel. +44 1223,496000, Fax: +44 1223496030

De Jong, J. HansWageningen Agricultural University, Graduate School of ExperimentalPlant Sciences, Laboratory of Genetics, Dreijenlaan 2, 6703 HAWageningen, The Netherlands, Tel: +31.317.482150, Fax: +31317483146;E-Mail: [email protected]

Dutrillaux, Bernard H.Institute Curie-CNRS, UMR 147,26 rue d' Ulm, F-75248 Paris Cedex 05;Departement de Radiobiologie et Radiopathologie, CENDSV, BP No 6,Fontenay-aux-Roses, France, Tel: +33 0142346672, Fax: +33 0142346674,E-Mail: [email protected]

Evans, D. Gareth R.Department of Medical Genetics, St Mary's Hospital, Hathersage Road,Manchester M13 OJH, UK

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List of Contributors XI

Federico, ConcettaDipartimento di Biologia Animale, University of Catania, via Androne 81,95124 Catania, Italy

FeU, R.Laboratory of Developmental Genetics and Imprinting, DevelopmentalGenetics Programme, The Brabraham Institute, Cambridge CB2 4AT, UK,Tel. +44 1223,496000, Fax: +44 1223496030

Ferguson-Smith, Malcolm A.Molecular Cytogenetics Group, University of Cambridge, Centre forVeterinary Science, Madingley Road, Cambridge CB3 OES, UK, Tel: +441223-333692, Fax: +44 1223-339067, E-Mail: [email protected]

Foresti, FaustoDepartamento de Morfologia, Instituto de Biociencias, UNESP, Campus deBotucatu, 18618-000, Botucatu, SP, Brasil

Franco, BrunellaTelethon Institute of Genetics and Medicine, San Raffaele BiomedicalScience Park, Via Olgettina 58,1-20132 Milano, Italy, Tel: +3902-21560­202, Fax: +3902-21560-220, E-Mail: [email protected]

Fransz, P. F.Laboratory of Molecular Biology, Wageningen Agricultural University,Graduate School of Experimental Plant Sciences, Dreijenlaan 2, NL-6703HAWageningen, The Netherlands

Fukui, KiichiDepartment of Biotechnology, Graduate School of Engineering, OsakaUniversity, Suita 565-0871, Osaka, Japan

Greaves, MartinCRC Section of Molecular Genetics, Paterson Institute for Cancer Research,Wilmslow Road, Manchester M20 4BX, UK

Hameister, HorstAbteilung Medizinische Genetik, Universitat Ulm, Albert-Einstein-Allee11, D-89069 Ulm, Germany, Tel: +49 731 5023436, Fax: +49 731 5023438,E-Mail: [email protected]

Harper, J. A.Institute of Grassland and Environmental Research, Plas Gogerddan,Aberystwyth, Ceredigion, SY23 3EB, Wales, UK

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XII List of Contributors

Heslop-Harrison, John S.Department of Cell Biology, John Innes Centre, Colney Lane, NorwichNR4 7UJ, England, Tel: +44 1603452571, Fax: +44 1603456844, E-Mail:[email protected]

Hong, Chia-sinDepartment of Biochemistry and Cellular and Molecular Biology,University of Tennessee, M407 Walters Life Sciences Bldg., Knoxville,Tennessee 37996-0840, USA, Tel: +1 423 974 3612, Fax: +1 4239746306

Ikemura, ToshimichiDepartment of Evolutionary Genetics, National Institute of Genetics, TheGraduate University for Advanced Studies, Mishima, Shizuoka-ken 411­8540 Japan, Tel: +81 55981 6788, Fax: +81 55981 6794, E-Mail: tikemu­[email protected]

Jacobsen, E.Laboratory of Plant Breeding, Wageningen Agricultural University,Graduate School of Experimental Plant Sciences, Dreijenlaan 2, NL-6703HAWageningen, The Netherlands

James, Louise A.CRC Section of Molecular Genetics, Paterson Institute for Cancer Research,Wilmslow Road, Manchester M20 4BX, UK

Jimenez, GuillermoDepartamento de Genetica, Facultad de Biologia, Universidad Complutense,E-28040 Madrid, Spain, Tel: +34 1 3945044, Fax: +341 3944844

Jones, R. N.Institute of Biological Sciences, University of Wales, Aberystwyth,Ceredigion, SY23 3EB, Wales, UK

Jones, T. A. H.Institute of Grassland and Environmental Research, PIas Gogerddan,Aberystwyth, Ceredigion, SY23 3EB, Wales, UK

Kanaya, ShigehikoFaculty of Engineering, Yamagata University, Yonezawa, Yamagata-ken992-8510, Japan

Kelsey, Anna M.Department of Histopathology, Royal Manchester Children's Hospital,Pendlebury, Manchester M27 4HA, UK

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List of Contributors XIII

Kelsey, G.Laboratory of Developmental Genetics and Imprinting, DevelopmentalGenetics Programme, The Brabraham Institute, Cambridge CB2 4AT, UK,Tel. +44 1223,496000, Fax: +44 1223496030

King, Ian P.Institute of Grassland and Environmental Research, Plas Gogerddan,Aberystwyth, Ceredigion, SY23 3EB, Wales, UK; Tel: +44 1970 828 255,Fax: +44 1970828 357; E-Mail: [email protected]

King, J.Institute of Grassland and Environmental Research, Plas Gogerddan,Aberystwyth, Ceredigion, SY23 3EB, Wales, UK

Kleckner, NancyDepartment of Molecular and Cellular Biology, Harvard University,Cambridge, MA 02138, USA, Tel: +I 617 495 4278, Fax: +1 617495 0758,E-Mail: [email protected]

Loidl, JosefInstitute of Botany, University of Vienna, Rennweg 14, A-1030 Vienna,Austria, Tel: +43 1 4277-54022, Fax: +43 1 4277-9541, E-Mail:[email protected]

Macgregor, HerbertDepartment of Biology, University of Leicester, University Road, LeicesterLEI 7RH, UK, Tel: +44 1162523344, Fax: +44116 252 3330, E-Mail:[email protected]

Maestra, BelenDepartamento de Genetica, Facultad de Biologia, UniversidadComplutense, E-28040 Madrid, Spain

Manzanero, SilviaDepartamento de Genetica, Facultad de Biologia, Universidad Complutense,28040 Madrid, Spain, Tel: +34 1 3945044, Fax: +34 1 3944844

McGown, GailCRC Section of Molecular Genetics, Paterson Institute for Cancer Research,Wilmslow Road, Manchester M20 4BX, UK

McKee, Bruce D.Department of Biochemistry and Cellular and Molecular Biology,University of Tennessee, M407 Walters Life Sciences Bldg., Knoxville,

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XIV List of Contributors

Tennessee 37996-0840, USA, Tel: +1 4239743612, Fax: +1423 9746306,E-Mail: [email protected]

Morgan, W. G.Institute of Grassland and Environmental Research, Plas Gogerddan,Aberystwyth, Ceredigion, SY23 3EB, Wales, UK

Murrell,A.Laboratory of Developmental Genetics and Imprinting, DevelopmentalGenetics Programme, The Babraham Institute, Cambridge CB2 4AT, UK

Naranjo, Carlos A.Instituto Fitotecnico Santa Catalina (FCAF, UNLP)-Centro deInvestigaciones Geneticas (UNLP-CONICET-CIC), C.CA 1836 Llavallol,Buenos Aires, Argentina

Naranjo, TomasDepartamento de Genetica, Facultad Biologia, Universidad Complutense,Ciudad Universitaria, E-28040 Madrid, Spain, Tel: +34 1 3945040, Fax:+34 1 3944844, E-Mail: [email protected]

Nash,J. V.Institute of Grassland and Environmental Research, Plas Gogerddan,Aberystwyth, Ceredigion, SY23 3EB, Wales, UK

Nicklas, R. BruceDepartments of Zoology and Cell Biology, Duke University, Box 91000,Durham, NC 27708-1000, USA, Tel: +1 919 6138196, Fax: +1 919 6138177, E-Mail: [email protected]

O'Brien, Patricia C. MMolecular Cytogenetics Group, University of Cambridge, Centre forVeterinary Science, Madingley Road, Cambridge CB3 OES, UK, Tel: +441223-333692, Fax: +44 1223-339067

Ohmido, NobukoLaboratory of Rice Genetic Engineering, Hokuriku National AgriculturalExperiment Station, Joetsu, 943-0193, Niigata, Japan

Ohno, MizukiDepartment of Evolutionary Genetics, National Institute of Genetics, TheGraduate University for Advanced Studies, Mishima, Shizuoka-ken 411­8540, Japan; Institute of Genetic Information, Kyushu University, Fukuoka812-0054, Japan

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List of Contributors xv

Pimpinelli, SergioIstituto Pasteur, Fondazione Cenci Bolognetti, Dipartimento di Genetica eBiologia Molecolare, Universita di Roma "La Sapienza", Piazzale AldoMoro 5,1-00185 Roma, Italy, Tel: +39649912876, Fax: +39644 56866,E-Mail: [email protected]

Poggio, LidiaDepartamento de Biologia, Facultad de Ciencias Exactas y Naturales,Universidad de Buenos Aires, 1428 Buenos Aires, Argentina

Puertas, Maria J.Departamento de Genetica, Facultad de Biologia, Universidad Complutense,E-28040 Madrid, Spain, Tel: +34 I 3945044, Fax: +34 1 3944844, E-Mail:[email protected]

Reik, WolfDevelopmental Genetics and Imprinting, Developmental GeneticsProgramme, The Babraham Institute, Cambridge CB2 4AT, UK, Tel: +441223496000, Fax: +44 1223496030, E-Mail: [email protected]

Rens, WillemMolecular Cytogenetics Group, University of Cambridge, Centre forVeterinary Science, Madingley Road, Cambridge CB3 OES, UK, Tel: +441223-333692, Fax: +44 1223-339067

Rettenberger, GuntherAbteilung Medizinische Genetik, Universitat VIm, Albert-Einstein-Allee11, D-89069 VIm, Germany; Labor Drs. Mehnert, Wegener Strasse 15,0-89231 Neu-Ulm, Germany

Roberts, L. A.Institute of Grassland and Environmental Research, Plas Gogerddan,Aberystwyth, Ceredigion, SY23 3EB, Wales, VK

Rosato, MarcelaDepartamento de Biologia, Facultad de Ciencias Exactas y Naturales,Universidad de Buenos Aires, 1428 Buenos Aires, Argentina

Saccone, SalvatoreDipartimento die Biologia Animale, University of Catania, via Androne 81,1-95124 Catania, Italy

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XVI List of Contributors

Taylor, SteveInstitute of Grassland and Environmental Research, Plas Gogerddan,Aberystwyth, Ceredigion, SY23 3EB, Wales, UK

Tenzen, ToyoakiDepartment of Evolutionary Genetics, National Institute of Genetics, TheGraduate University for Advanced Studies, Mishima, Shizuoka-ken 411­8540 Japan, Tel: +81 55981 6788, Fax: +81 55981 6794

Thomas, H. M.Institute of Grassland and Environmental Research, Plas Gogerddan,Aberystwyth, Ceredigion, SY23 3EB, Wales, UK

Thorncroft, MaryCRC Section of Molecular Genetics, Paterson Institute for Cancer Research,Wilmslow Road, Manchester M20 4BX, UK

Varley, Jennifer M.CRC Section of Molecular Genetics, Paterson Institute for CancerResearch, Wilmslow Road, Manchester M20 4BX, UK, Tel: +44 161 4463062, Fax: +44 1614463062, E-Mail: [email protected]

Vogel, WaltherAbteilung Medizinische Genetik, Universitat Ulm, Albert-Einstein-Allee11, D-89069 Ulm, Germany

Wako, ToshiyukiLaboratory of Rice Genetic Engineering, Hokuriku National AgriculturalExperiment Station, Joetsu, 943-0193, Niigata, Japan

Walter, J.Max-Planck Institute of Molecular Genetics, Berlin, Germany

Watanabe, YoshihisaDepartment of Evolutionary Genetics, National Institute of Genetics, TheGraduate University for Advanced Studies, Mishima, Shizuoka-ken 411­8540 Japan, Tel: +81 55981 6788, Fax: +81 55981 6794

Wennekes-van Eden, J.Laboratory of Plant Breeding, Wageningen Agricultural University,Graduate School of Experimental Plant Sciences, Dreijenlaan 2, 6703HAWageningen, The Netherlands

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List of Contributors XVII

Yamagata, TesushiDepartment of Evolutionary Genetics, National Institute of Genetics, TheGraduate University for Advanced Studies, Mishima, Shizuoka-ken 411­8540 Japan, Tel: +81 55981 6788, Fax: +81 55981 6794

Yang, FengtangMolecular Cytogenetics Group, University of Cambridge, Centre forVeterinary Science, Madingley Road, Cambridge CB3 OES, UK, Tel: +441223-333692, Fax: +44 1223-339067

Yoo, SiukDepartment of Biochemistry and Cellular and Molecular Biology,University of Tennessee, M407 Walters Life Sciences Bldg., Knoxville,Tennessee 37996-0840, USA, Tel: +1 4239743612, Fax: +1 4239746306

Zabel, P.Laboratory of Molecular Biology, Wageningen Agricultural University,Graduate School of Experimental Plant Sciences, Dreijenlaan 2, NL-6703HAWageningen, The Netherlands

Zhong, X.-B.Yale University, School of Medicine 1-310 SHM, New Haven, CT 06520­8005, USA

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XXI

Dedication

In the months that have preceded and followed the 13th Conference, three out­standing Italian cytogeneticists have disappeared.

Mario Benazzi, a pioneer of Evolutionary Cytogenetics, died in December1997 after a long and fruitful life mainly devoted to the study of Plathelmintescytogenetics.

Alessandro Morescalchi, the outstanding and respected student ofAmphibiancytogenetics, with whom E. Olmo shared work and friendship for many years,died untimely in March 1997.

Marco Milani-Comparetti, an outstanding medical cytogeneticist and expertin bioethics, a member of the local Organising Committee, died in May 1999.

We wish to dedicate to them this 13th volume of Chromosomes Today.

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XXIII

Preface

The 13th International Chromosome Conference that was held in Ancona andNumana from 8th to 12th September 1998 was the last of this century andoffered a unique opportunity to draw a balance of more than one hundred yearsof chromosome research.

The participation of over 300 researchers and the high scientific quality ofthe contributions represent a significant index of the important results obtainedso far and of the ever growing interest raised by cytogenetic studies. This alsohelped us to understand the importance of the insight that led C.D. Darlingtonto found, in Oxford in 1964, what developed into the InternationalChromosome Conference so that scientists working on different aspects ofchromosome research would have the opportunity of "discussing the funda­mental problems of chromosomes with one another".

The present volume of the series Chromosomes Today contains all but oneof the invited lectures given at the 13th International ChromosomeConference. The programme, the poster presentations, the poster abstracts andthe full list of the organisers and of the members of the Advisory Committeehave been published in Cytogenetics and Cell Genetics vol. 81 (2): 91-168,September 1998.

In the previous volume of the series it was observed that molecular biologyhas become definitively incorporated into cytogenetics. The articles collectedin this volume are a witness to the fruitfulness of this integration.

A new, important aspect that emerged from the 13th InternationalChromosome Conference are the advances made in the new, powerful meth­ods for studying chromosomes derived from the amazing synergy of molecu­lar biology, optical science and computing technology. However, the importantmethodological developments and the new applicative opportunities that haveoccurred in our field have not diminished the attention towards some essentialissues such as the composition and organisation of the interphase nucleus, theinteraction of homologous chromosomes in meiotic events, the regulation ofchromosome distribution during division, or the role of genome structural andmolecular variations during evolution. The relevant recent advances in allthese issues are represented and discussed in this volume of ChromosomesToday.

The organisation of the 13th International Chromosome Conference hasbeen a demanding undertaking repaid by great satisfaction. For their hard workand their commitment we are grateful to the Standing Committee, theInternational Scientific Advisory Committee, the local Organising Committee,

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XXIV Preface

many volunteer helpers, the University of Ancona and several other bodieswhich provided both financial and organisational support.

The 14th International Chromosome Conference will be held at WuerzburgUniversity, Germany, under the guidance of Michael Schmid. We are lookingforward to the further advances that will have been made in chromosome stud­ies at the beginning of the third millennium and hope that the InternationalChromosome Conference will continue to be one of the most important oppor­tunities for scientists from the different fields of cytogenetics to meet and todiscuss and exchange data.

The Editors