march 2014
genesis The Journal of Genetics and Development Special Issue on Sea Urchins
ISSN 1526-968X
volume 52 no. 3
editor sally a. moody guest editors: Charles Ettensohn and David McClay
Editor-in-Chief Sally A. Moody George Washington University 207 Ross Hall, 2300 I Street, NW Washington DC 20037 Tel: 202-994-2878 E-mail: [email protected]
Associate Editors Deborah L. Chapman Department of Biological Sciences University of Pittsburgh Room 141 Life Sciences Annex 4249 Fifth Avenue Pittsburgh, PA 15260 Tel: 412-624-0774 Lab: 412-624-0580 Fax: 412-624-4759 E-mail: [email protected]
Kathryn Cheah Department of Biochemistry University of Hong Kong Sassoon Rd Hong Kong SAR China Tel: 852-2819-9240 Fax: 852-2855-1254 E-mail: [email protected]
Justin Kumar Department of Biology Indiana University Jordan Hall A318 1001 East Third Street Bloomington, IN 47405 Tel: 812-856-2621 Fax: 812-856-1566 E-mail: [email protected] Website: www.bio.indiana. edu/~kumarlab
José Xavier-Neto LNBio Campus do LNLS: Rua Giuseppe Máximo Scolfaro, 10000 Pólo Il de Alta Tecnologia de Campinas Campinas, SP CEP 13083-970 Brasil Tel: +55 19 3512-1249 Fax: +55 19 3512-1004 E-mail: [email protected]
Valerie Reinke Department of Genetics Yale University School of Medicine PO Box 208005 233 Cedar Street New Haven CT 06520-8005 Tel: 203-785-5228 Fax: 203-785-6350 E-mail: [email protected]
Editorial Board Enrique Amaya Manchester, United Kingdom Chris T. Amemiya Seattle, Washington Helena Araujo Rio de Janeiro, RJ, Brazil Kavita Arora Irvine, Calfornia Thomas Becker Camperdown, Australia Richard Behringer Houston, Texas Rolf Bodmer La Jolla, California Véronique Borday-Birraux Gif-sur-Yvette, France Maja Bucan Philadelphia, Pennsylvania Kwang-Wook Choi Daejeon, Korea
Bon-chu Chung Taipei, Taiwan Sophie Creuzet Gif-sur-Yvette, France Liam Dolan Norwich, United Kingdom Charles A. Ettensohn Pittsburgh, Pennsylvania Peter Farlie Parkville Victoria, Australia Bernd Fritzsch Iowa City, Iowa Xiang Gao Nanjing, China Tom Gridley Bar Harbor, Maine Anna-Katerina Hadjantonakis New York, New York Marnie E. Halpern Baltimore, Maryland
Ian Hope Leeds, United Kingdom
Terry Magnuson Chapel Hill, North Carolina
Andrew Hudson Edinburgh, United Kingdom
William Mattox Houston, Texas
Randy Johnson Houston, Texas
David R. McClay Durham, North Carolina
Susan Kidson Cape Town, South Africa
David M. Miller, III Nashville, Tennessee
James Kramer Chicago, lIIinois
Yuji Mishina Ann Arbor, Michigan
Tsutomu Kume Chicago, lIIinois
Mary Mullins Philadelphia, Pennsylvania
Michel Labouesse IIIkirch, France
Andras Nagy Toronto, Canada
Patrick Lemaire Marseille, France
Petra Pandur Ulm, Germany
Susan Mackem Bethesda, Maryland
Helen Salz Cleveland, Ohio
Yoshiki Sasai Kobe,Japan Nori Satoh Uruma, Japan Michael Scanlon Ithaca, New York John Schimenti Ithaca, New York Guojun Sheng Kobe, Japan Patrick Tam Wentworthville, Australia Gerald Thomsen Stony Brook, New York Christopher Wright Nashville, Tennessee Lyle Zimmerman London, United Kingdom Mario Zurita Cuernavaca Morelos, México
Disclaimer. The Publisher and Editors cannot be held responsible for errors or any consequences arising from the use of information contained in this journal; the views and opinions expressed do not necessarily reflect those of the Publisher and Editors, neither does the publication of advertisements constitute any endorsement by the Publisher and Editors of the products advertised. Copyright and copying. Copyright© 2014 Wiley Periodicals, Inc. All rights reserved. No part of this publication may be reproduced, stored or transmitted in any form or by any means without the prior permission in writing from the copyright holder. Authorization to copy items for internal and personal use is granted by the copyright holder for libraries and other users registered with their local Reproduction Rights Organisation (RRO), e.g. Copyright Clearance Center (CCC), 222 Rosewood Drive, Danvers, MA 01923, USA (www.copyright.com), provided the appropriate fee is paid directly to the RRO. This consent does not extend to other kinds of copying such as copying for general distribution, for advertising or promotional purposes, for creating new collective works or for resale. Special requests should be addressed to: [email protected] Information for subscribers. genesis is published in 12 issues per year. Institutional subscription prices for 2014 are: Online: US$3,543 (US), US$3,543 (and Rest of World), €2,289 (Europe), £1,809 (UK). Prices are exclusive of tax. Asia-Pacific GST, Canadian GST and European VAT will be applied at the appropriate rates. For more information on current tax rates, please go to www.wileyonlinelibrary.com/tax-vat. The price includes online access to the current and all online back files to January 1st 2008, where available. For other pricing options, including access information and terms and conditions, please visit www.wileyonlinelibrary.com/access. Back issues. Single issues from current and recent volumes are available at the current single issue price from [email protected]. Earlier issues may be obtained from Periodicals Service Company, 11 Main Street, Germantown, NY 12526, USA. Tel: +1 518 537 4700, Fax: +1 518 537 5899, Email: [email protected] Journal Customer Services. For ordering information, claims and any enquiry concerning your journal subscription please go to www.wileycustomerhelp.com/ask or contact your nearest office. Americas: Email: [email protected]; Tel: +1 781 388 8598 or +1 800 835 6770 (toll free in the USA & Canada). Europe, Middle East and Africa: Email: [email protected]; Tel: +44 (0) 1865 778315. Asia Pacific: Email: [email protected]; Tel: +65 6511 8000. Japan: For Japanese speaking support, Email: [email protected]; Tel: +65 6511 8010 or Tel (toll-free): 005 316 50 480. Visit our Online Customer Get-Help available in 6 languages at www.wileycustomerhelp.com Access to this journal is available free online within institutions in the developing world through the AGORA initiative with the FAO, the HINARI initiative with the WHO and the OARE initiative with UNEP. For information, visit www.aginternetwork.org, www.healthinternetwork.org, www.oaresciences.org Wiley’s Corporate Citizenship initiative seeks to address the environmental, social, economic, and ethical challenges faced in our business and which are important to our diverse stakeholder groups. Since launching the initiative, we have focused on sharing our content with those in need, enhancing community philanthropy, reducing our carbon impact, creating global guidelines and best practices for paper use, establishing a vendor code of ethics, and engaging our colleagues and other stakeholders in our efforts. Follow our progress at www.wiley.com/go/citizenship Aims and Scope. As of January 2000, Developmental Genetics was renamed and relaunched as genesis: The Journal of Genetics and Development, with a new scope and Editorial Board. The journal focuses on work that addresses the genetics of development and the fundamental mechanisms of embryological processes in animals and plants. With increased awareness of the interplay between genetics and evolutionary change, particularly during developmental processes, we encourage submission of manuscripts from all ecological niches. The expanded numbers of genomes for which sequencing is being completed will facilitate genetic and genomic examination of developmental issues, even if the model system does not fit the “classical genetic” mold. Therefore, we encourage submission of manuscripts from all species. Other areas of particular interest include: 1) the roles of epigenetics, microRNAs and environment on developmental processes; 2) genome-wide studies; 3) novel imaging techniques for the study of gene expression and cellular function; 4) comparative genetics and genomics and 5) animal models of human genetic and developmental disorders. genesis presents reviews, full research articles, short research letters, and state-of-the-art technology reports that promote an understanding of the function of genes and the roles they play in complex developmental processes. genesis accepts articles for Open Access publication. Please visit http://olabout.wiley.com/WileyCDA/Section/id-406241.html for further information about OnlineOpen. Cover illustration: A pluteus larva of the sand dollar Mellita quinquiesperforatus. The skeleton was stained with calcein and the nuclei were stained with Hoechst stain. Shown is a complete stack of confocal images. Image by Dede Lyons, Julian Kimura and Megan Martik of the McClay lab. ISSN 1526-968X (Online)
genesis The Journal of Genetics and Development Special Issue on Sea Urchins
ISSN 1526-968X
volume 52 no. 3
editor sally a. moody guest editors: Charles Ettensohn and David McClay
Editor-in-Chief Sally A. Moody George Washington University 207 Ross Hall, 2300 I Street, NW Washington DC 20037 Tel: 202-994-2878 E-mail: [email protected]
Associate Editors Deborah L. Chapman Department of Biological Sciences University of Pittsburgh Room 141 Life Sciences Annex 4249 Fifth Avenue Pittsburgh, PA 15260 Tel: 412-624-0774 Lab: 412-624-0580 Fax: 412-624-4759 E-mail: [email protected]
Kathryn Cheah Department of Biochemistry University of Hong Kong Sassoon Rd Hong Kong SAR China Tel: 852-2819-9240 Fax: 852-2855-1254 E-mail: [email protected]
Justin Kumar Department of Biology Indiana University Jordan Hall A318 1001 East Third Street Bloomington, IN 47405 Tel: 812-856-2621 Fax: 812-856-1566 E-mail: [email protected] Website: www.bio.indiana. edu/~kumarlab
José Xavier-Neto LNBio Campus do LNLS: Rua Giuseppe Máximo Scolfaro, 10000 Pólo Il de Alta Tecnologia de Campinas Campinas, SP CEP 13083-970 Brasil Tel: +55 19 3512-1249 Fax: +55 19 3512-1004 E-mail: [email protected]
Valerie Reinke Department of Genetics Yale University School of Medicine PO Box 208005 233 Cedar Street New Haven CT 06520-8005 Tel: 203-785-5228 Fax: 203-785-6350 E-mail: [email protected]
Editorial Board Enrique Amaya Manchester, United Kingdom Chris T. Amemiya Seattle, Washington Helena Araujo Rio de Janeiro, RJ, Brazil Kavita Arora Irvine, Calfornia Thomas Becker Camperdown, Australia Richard Behringer Houston, Texas Rolf Bodmer La Jolla, California Véronique Borday-Birraux Gif-sur-Yvette, France Maja Bucan Philadelphia, Pennsylvania Kwang-Wook Choi Daejeon, Korea
Bon-chu Chung Taipei, Taiwan Sophie Creuzet Gif-sur-Yvette, France Liam Dolan Norwich, United Kingdom Charles A. Ettensohn Pittsburgh, Pennsylvania Peter Farlie Parkville Victoria, Australia Bernd Fritzsch Iowa City, Iowa Xiang Gao Nanjing, China Tom Gridley Bar Harbor, Maine Anna-Katerina Hadjantonakis New York, New York Marnie E. Halpern Baltimore, Maryland
Ian Hope Leeds, United Kingdom
Terry Magnuson Chapel Hill, North Carolina
Andrew Hudson Edinburgh, United Kingdom
William Mattox Houston, Texas
Randy Johnson Houston, Texas
David R. McClay Durham, North Carolina
Susan Kidson Cape Town, South Africa
David M. Miller, III Nashville, Tennessee
James Kramer Chicago, lIIinois
Yuji Mishina Ann Arbor, Michigan
Tsutomu Kume Chicago, lIIinois
Mary Mullins Philadelphia, Pennsylvania
Michel Labouesse IIIkirch, France
Andras Nagy Toronto, Canada
Patrick Lemaire Marseille, France
Petra Pandur Ulm, Germany
Susan Mackem Bethesda, Maryland
Helen Salz Cleveland, Ohio
Yoshiki Sasai Kobe,Japan Nori Satoh Uruma, Japan Michael Scanlon Ithaca, New York John Schimenti Ithaca, New York Guojun Sheng Kobe, Japan Patrick Tam Wentworthville, Australia Gerald Thomsen Stony Brook, New York Christopher Wright Nashville, Tennessee Lyle Zimmerman London, United Kingdom Mario Zurita Cuernavaca Morelos, México
Disclaimer. The Publisher and Editors cannot be held responsible for errors or any consequences arising from the use of information contained in this journal; the views and opinions expressed do not necessarily reflect those of the Publisher and Editors, neither does the publication of advertisements constitute any endorsement by the Publisher and Editors of the products advertised. Copyright and copying. Copyright© 2014 Wiley Periodicals, Inc. All rights reserved. No part of this publication may be reproduced, stored or transmitted in any form or by any means without the prior permission in writing from the copyright holder. Authorization to copy items for internal and personal use is granted by the copyright holder for libraries and other users registered with their local Reproduction Rights Organisation (RRO), e.g. Copyright Clearance Center (CCC), 222 Rosewood Drive, Danvers, MA 01923, USA (www.copyright.com), provided the appropriate fee is paid directly to the RRO. This consent does not extend to other kinds of copying such as copying for general distribution, for advertising or promotional purposes, for creating new collective works or for resale. Special requests should be addressed to: [email protected] Information for subscribers. genesis is published in 12 issues per year. Institutional subscription prices for 2014 are: Online: US$3,543 (US), US$3,543 (and Rest of World), €2,289 (Europe), £1,809 (UK). Prices are exclusive of tax. Asia-Pacific GST, Canadian GST and European VAT will be applied at the appropriate rates. For more information on current tax rates, please go to www.wileyonlinelibrary.com/tax-vat. The price includes online access to the current and all online back files to January 1st 2008, where available. For other pricing options, including access information and terms and conditions, please visit www.wileyonlinelibrary.com/access. Back issues. Single issues from current and recent volumes are available at the current single issue price from [email protected]. Earlier issues may be obtained from Periodicals Service Company, 11 Main Street, Germantown, NY 12526, USA. Tel: +1 518 537 4700, Fax: +1 518 537 5899, Email: [email protected] Journal Customer Services. For ordering information, claims and any enquiry concerning your journal subscription please go to www.wileycustomerhelp.com/ask or contact your nearest office. Americas: Email: [email protected]; Tel: +1 781 388 8598 or +1 800 835 6770 (toll free in the USA & Canada). Europe, Middle East and Africa: Email: [email protected]; Tel: +44 (0) 1865 778315. Asia Pacific: Email: [email protected]; Tel: +65 6511 8000. Japan: For Japanese speaking support, Email: [email protected]; Tel: +65 6511 8010 or Tel (toll-free): 005 316 50 480. Visit our Online Customer Get-Help available in 6 languages at www.wileycustomerhelp.com Access to this journal is available free online within institutions in the developing world through the AGORA initiative with the FAO, the HINARI initiative with the WHO and the OARE initiative with UNEP. For information, visit www.aginternetwork.org, www.healthinternetwork.org, www.oaresciences.org Wiley’s Corporate Citizenship initiative seeks to address the environmental, social, economic, and ethical challenges faced in our business and which are important to our diverse stakeholder groups. Since launching the initiative, we have focused on sharing our content with those in need, enhancing community philanthropy, reducing our carbon impact, creating global guidelines and best practices for paper use, establishing a vendor code of ethics, and engaging our colleagues and other stakeholders in our efforts. Follow our progress at www.wiley.com/go/citizenship Aims and Scope. As of January 2000, Developmental Genetics was renamed and relaunched as genesis: The Journal of Genetics and Development, with a new scope and Editorial Board. The journal focuses on work that addresses the genetics of development and the fundamental mechanisms of embryological processes in animals and plants. With increased awareness of the interplay between genetics and evolutionary change, particularly during developmental processes, we encourage submission of manuscripts from all ecological niches. The expanded numbers of genomes for which sequencing is being completed will facilitate genetic and genomic examination of developmental issues, even if the model system does not fit the “classical genetic” mold. Therefore, we encourage submission of manuscripts from all species. Other areas of particular interest include: 1) the roles of epigenetics, microRNAs and environment on developmental processes; 2) genome-wide studies; 3) novel imaging techniques for the study of gene expression and cellular function; 4) comparative genetics and genomics and 5) animal models of human genetic and developmental disorders. genesis presents reviews, full research articles, short research letters, and state-of-the-art technology reports that promote an understanding of the function of genes and the roles they play in complex developmental processes. genesis accepts articles for Open Access publication. Please visit http://olabout.wiley.com/WileyCDA/Section/id-406241.html for further information about OnlineOpen. Cover illustration: A pluteus larva of the sand dollar Mellita quinquiesperforatus. The skeleton was stained with calcein and the nuclei were stained with Hoechst stain. Shown is a complete stack of confocal images. Image by Dede Lyons, Julian Kimura and Megan Martik of the McClay lab. ISSN 1526-968X (Online)
