Desarrollo embrionario Keith L.

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& T.V.N. Persaud & Mark G. Torchia
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KEITH L. MOORE
El Dr. Moore ha obtenido numerosos premios y reconocimientos de
prestigio. Ha recibido las máximas condecoraciones por su
destacado historial de publicaciones de libros de anatomía y
embriología con orientación clínica. Fue galardonado con el primer
Henry Gray/Elsevier Distinguished Educator Award en 2007, la
máxima condecoración otorgada por la American Association of
Anatomists en reconocimiento a la excelencia en la enseñanza de la
anatomía humana en estudios de grado y doctorado de ciencias
médicas y odontológicas; galardonado también con el Honored
Member Award de la American Association of Clinical Anatomists
(1994) por sus notables contribuciones en el campo de la anatomía
clínica, y con el J.C.B. Grant Award de la Canadian Association of
Anatomists (1984) «en reconocimiento a su meritorio servicio y a su
extraordinaria erudición en el campo de las ciencias anatómicas». En
2008, el profesor Moore pasó a ser Fellow de la American
Association of Anatomists (AAA). El rango de Fellow honra a los
miembros distinguidos de la AAA que han alcanzado cotas de
excelencia en su desarrollo científico y en sus contribuciones a las
ciencias médicas. En 2012, el Dr. Moore recibió el grado de Honorary
Doctor of Science por la Ohio State University y por la University of
Western Ontario en 2015; la Queen Elizabeth II Diamond Jubilee
Medal canadiense en honor de sus notables contribuciones y logros,
y el Benton Adkins Jr. Distinguished Service Award por su
extraordinaria hoja de servicios a la American Association of Clinical
Anatomists.

T.V.N. (VID) PERSAUD

El Dr. Persaud fue galardonado con el Henry Gray/Elsevier
Distinguished Educator Award en 2010, «la máxima distinción de la
American Association of Anatomists en reconocimiento a la
excelencia continuada y el liderazgo en la enseñanza de la anatomía
humana»; con el Honored Member Award de la American
Association of Clinical Anatomists (2008) por «su distinguida
carrera y sus notables contribuciones en el campo de la anatomía
clínica, la embriología y la historia de la anatomía, y con el J.C.B.
Grant Award de la Canadian Association of Anatomists (1991) «en
reconocimiento a su meritorio servicio y a su extraordinaria
erudición en el campo de las ciencias anatómicas». En 2010, el
profesor Persaud pasó a ser Fellow de la American Association of
Anatomists. El rango de Fellow honra a los miembros distinguidos
de la AAA que han alcanzado cotas de excelencia en su desarrollo
científico y en sus contribuciones a las ciencias médicas. En 2003, el
Dr. Persaud fue galardonado con la Queen Elizabeth II Golden
Jubilee Medal, nominado por el Gobierno de Canadá, por «su
notable contribución a la nación, a la comunidad y a sus
compatriotas canadienses».

MARK G. TORCHIA
El Dr. Mark G. Torchia recibió el primer Governor General Award
for Innovation, que «reconoce y celebra a las personas, equipos y
organizaciones canadienses destacados, pioneros y creadores que
contribuyen al éxito de nuestro país, que ayudan a configurar
nuestro futuro y que inspiran a la siguiente generación». El Dr.
Torchia también ha recibido el Manning Principle Prize (2015), que
reconoce a los «líderes y visionarios que tienen un impacto positivo
en la economía canadiense a la vez que mejoran la experiencia
humana en todas sus dimensiones alrededor del mundo».
Asimismo, ha recibido el Norman and Marion Bright Memorial
Medal and Award para «los individuos que han realizado una
contribución destacada a la tecnología química» y el TIMEC
Medical Device Champion Award. El Dr. Torchia sigue implicado
con estudiantes de todos los niveles mediante actividades de
divulgación e impartición de cursos. Ha sido nominado para los
premios a la docencia de la Manitoba Medical Students’ Association
(MMSA) desde su inicio y ha sido galardonado con el Award for
Teaching Excellence (2016) de la Rady Faculty of Health Sciences,
University of Manitoba.
Desarrollo embrionario

DÉCIMA EDICIÓN

Keith L. Moore, BA, MSc, PhD, DSc

(OSU), DSc (WU), FIAC, FRSM, FAAA
Professor Emeritus, Division of Anatomy, Department of Surgery
Former Professor and Chair, Department of Anatomy, and Associate Dean
for Basic Medical Sciences
Faculty of Medicine, University of Toronto, Toronto, Ontario, Canada
Former Professor and Head of Anatomy, Faculty of Medicine, University of
Manitoba, Winnipeg, Manitoba, Canada

T.V.N. (Vid) Persaud, MD, PhD, DSc,

FRCPath (Lond.), FAAA
Professor Emeritus and Former Head, Department of Human Anatomy and
Cell Science
Professor of Pediatrics and Child Health
Associate Professor of Obstetrics, Gynecology, and Reproductive Sciences,
Max Rady College of Medicine, Faculty of Health Sciences, Faculty of
Medicine, University of Manitoba, Winnipeg, Manitoba, Canada
Part-Time Professor of Anatomy, St. George’s University, Grenada, West
Indies
Mark G. Torchia, MSc, PhD
Associate Professor, Department of Surgery; Associate Professor,
Department of Human Anatomy and Cell Sciences, Max Rady College of
Medicine, Rady Faculty of Health Sciences
Executive Director, Centre for the Advancement of Teaching and Learning,
Vice-Provost (Teaching and Learning) University of Manitoba, Winnipeg,
Manitoba, Canada

Acceda al contenido electrónico en inglés en

StudentConsult.com
Índice de capítulos

Instrucciones para el acceso en línea

Cubierta

Portada

Página de créditos

Dedicatoria

Colaboradores

Prefacio

Agradecimientos

Saca el mayor partido de la 10.ª edición de Desarrollo embrionario.

1: Introducción al desarrollo humano

Importancia y avances en embriología

 Términos descriptivos

2: Reproducción humana

Órganos reproductores

Gametogénesis

Ciclo reproductivo femenino

Transporte de los gametos

Maduración de los espermatozoides

Viabilidad de los ovocitos y los espermatozoides

3: Primera semana del desarrollo humano

Fecundación

Segmentación del cigoto

Formación del blastocisto

4: Segunda semana del desarrollo humano

Formación de la cavidad amniótica, el disco embrionario y la

vesícula umbilical

Desarrollo del saco coriónico

Lugares de implantación de los blastocistos

5: Tercera semana del desarrollo humano

 Gastrulación: formación de las capas germinativas

Neurulación: formación del tubo neural

Desarrollo de los somitas

Desarrollo del celoma intraembrionario

Desarrollo temprano del sistema cardiovascular

Desarrollo de las vellosidades coriónicas

6: Cuarta a octava semanas del desarrollo humano

Plegamiento del embrión

Derivados de las capas germinativas

Control del desarrollo embrionario

Estimación de la edad embrionaria

Aspectos destacados de las semanas cuarta a octava

7: Periodo fetal: de la novena semana al nacimiento

Aspectos destacados del periodo fetal

Factores que influyen en el crecimiento fetal

Procedimientos para evaluar la situación fetal

Periodo neonatal

8: Placenta y membranas fetales

 Placenta

Membrana amniocoriónica

Parto

Amnios y líquido amniótico

Vesícula umbilical

Alantoides

Embarazos múltiples

9: Cavidades corporales, mesenterios y diafragma

Cavidad corporal embrionaria

Desarrollo del diafragma

10: Aparato faríngeo, cara y cuello

Arcos faríngeos

Bolsas faríngeas

Hendiduras faríngeas

Membranas faríngeas

Desarrollo de la glándula tiroides

Desarrollo de la lengua

Desarrollo de las glándulas salivares

Desarrollo de la cara
 Desarrollo de las cavidades nasales

Desarrollo del paladar

11: Aparato respiratorio

Desarrollo de la laringe

Desarrollo de la tráquea

Desarrollo de los bronquios y los pulmones

12: Aparato digestivo

Intestino primitivo anterior

Desarrollo del bazo

Intestino primitivo medio

Intestino primitivo posterior

Sistema nervioso entérico

13: Aparato urogenital

Desarrollo del aparato urinario

Desarrollo de las glándulas suprarrenales

Desarrollo del aparato genital

Desarrollo de los conductos inguinales

Recolocación de los testículos y los ovarios

14: Aparato cardiovascular

Desarrollo temprano del corazón y los vasos sanguíneos

Desarrollo tardío del corazón

Anomalías congénitas del corazón y de los grandes vasos

Derivados de las arterias de los arcos faríngeos

Anomalías congénitas de las arterias de los arcos faríngeos

Circulación fetal y neonatal

Desarrollo del sistema linfático

15: Sistema musculoesquelético

Sistema esquelético

Desarrollo del cartílago y del hueso

Desarrollo de las articulaciones

Desarrollo del esqueleto axial

Desarrollo del esqueleto apendicular

Sistema muscular

Desarrollo de los miembros

16: Sistema nervioso

Desarrollo del sistema nervioso

Desarrollo de la médula espinal

 Anomalías congénitas de la médula espinal

Desarrollo del encéfalo

Anomalías congénitas del encéfalo

Desarrollo del sistema nervioso periférico

Desarrollo del sistema nervioso autónomo

17: Desarrollo de los ojos y los oídos

Desarrollo de los ojos y estructuras relacionadas

Desarrollo de los oídos

18: Sistema tegumentario

Desarrollo de la piel y sus anexos

Desarrollo del pelo

Desarrollo de las uñas

Desarrollo de las glándulas mamarias

Desarrollo de los dientes

19: Anomalías congénitas humanas

Teratología: estudio del desarrollo patológico

Anomalías congénitas causadas por factores genéticos*

Anomalías congénitas causadas por factores ambientales

 Anomalías congénitas causadas por herencia multifactorial

20: Bases celulares y moleculares del desarrollo

Comunicación intercelular

Morfógenos

Receptores tirosina cinasa

Vía Notch-Delta

Factores de transcripción

Epigenética

Células madre: diferenciación versus pluripotencialidad

Edición de genes

Resumen de las vías habituales de señalización usadas durante

el desarrollo

Bibliografía y lecturas recomendadas

Apéndice: respuestas a las preguntas orientadas clínicamente

Índice alfabético
Página de créditos

Avda. Josep Tarradellas, 20-30, 1.°, 08029, Barcelona, España

Before We Are Born: Essentials of Embryology and Birth Defects,

10th edition
Copyright © 2020, Elsevier Inc. All rights reserved.
Previous editions copyrighted 2016, 2013, 2008, 2003, 1998, 1993,
1989, 1983 and 1974
ISBN: 978-0-323-60849-7

This translation of Before We Are Born: Essentials of Embryology and

Birth Defects, 10th ed., by Keith L. Moore, T.V.N. (Vid) Persaud and
Mark G. Torchia, was undertaken by Elsevier España, S.L.U. and is
published by arrangement with Elsevier, Inc.

Esta traducción de Before We Are Born: Essentials of Embryology and

Birth Defects, 10.ª ed., de Keith L. Moore, T.V.N. (Vid) Persaud y Mark
G. Torchia, ha sido llevada a cabo por Elsevier España, S.L.U. y se
publica con el permiso de Elsevier, Inc.

Desarrollo embrionario, 10.ª ed., de Keith L. Moore, T.V.N. (Vid)

Persaud y Mark G. Torchia
© 2021 Elsevier España, S.L.U.
ISBN: 978-84-9113-958-4
eISBN: 978-84-1382-069-9

Todos los derechos reservados.

Reserva de derechos de libros

Cualquier forma de reproducción, distribución, comunicación
pública o transformación de esta obra solo puede ser realizada con la
autorización de sus titulares, salvo excepción prevista por la ley.
Diríjase a CEDRO (Centro Español de Derechos Reprográficos) si
necesita fotocopiar o escanear algún fragmento de esta obra
(www.conlicencia.com; 91 702 19 70/93 272 04 45).

A d v e rt en ci a
Esta traducción ha sido llevada a cabo por Elsevier España, S.L.U.
bajo su única responsabilidad. Facultativos e investigadores deben
siempre contrastar con su propia experiencia y conocimientos el
uso de cualquier información, método, compuesto o experimento
descrito aquí. Los rápidos avances en medicina requieren que los
diagnósticos y las dosis de fármacos recomendadas sean siempre
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asumen responsabilidad alguna por la traducción ni por los daños
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productos defectuosos o negligencia, o como consecuencia de la
aplicación de métodos, productos, instrucciones o ideas contenidos
en esta obra.

Revisión científica:
Concepción Martínez Álvarez
Catedrática de Universidad
Departamento de Anatomía y Embriología
Facultad de Medicina
Universidad Complutense de Madrid

Servicios editoriales: DRK Edición

Depósito legal: B 6899-2021
Impreso en España
Dedicatoria

En recuerdo de Marion

A mi amada esposa y mi mejor amiga, por su apoyo y paciencia

infinitos. Agradezco su aliento y comprensión durante las
incontables horas dedicadas a escribir las primeras tres ediciones de
Desarrollo embrionario. Mis maravillosos recuerdos la mantienen
viva en mi corazón y mi mente. Agradezco el continuo apoyo que he
recibido de mis hijas Pam y Kate y quiero expresar mi gratitud a mi
yerno, Ron Crowe, por su capacidad técnica. Estoy muy orgulloso de
mis cinco hijos, Warren, Pam, Karen, Laurel y Kate, de nuestros
nueve nietos, Kristin, Lauren, Caitlin, Mitchel, Jayme, Courtney,
Brooke, Melissa y Alicia, así como de nuestros bisnietos, James y
Charlotte.

-KLM

Para Gisela

Mi amada esposa y mi mejor amiga, por su apoyo y paciencia

infinitos; a nuestros tres hijos, Indrani, Sunita y Rainer (Ren), y
nuestros nietos (Brian, Amy y Lucas).

-TVNP

Para Barbara, Muriel y Erik

Gracias por vuestro apoyo, aliento, risas y amor. Vuestros logros

personales siguen asombrándome. Este libro está dedicado a
 vosotros.

-MGT

Para los alumnos y sus profesores

A nuestros alumnos: esperamos que disfrutéis con la lectura de este

libro, que amplíe vuestros conocimientos sobre embriología humana,
que aprobéis todos vuestros exámenes y que os sintáis emocionados
y bien preparados cuando tengáis que atender a vuestros pacientes,
así como cuando os apliquéis en tareas de investigación y de
docencia. Os quedaréis con algo de lo que escuchéis, gran parte de lo
que leáis, una parte aún mayor de lo que veáis y con casi todo lo que
experimentéis y comprendáis completamente. A sus profesores:
deseamos que este libro constituya un recurso útil para vosotros y
para vuestros alumnos. Apreciamos los numerosos y constructivos
comentarios que hemos recibido a lo largo de los años, tanto de
alumnos como de profesores. Vuestras observaciones han sido
inestimables para que hayamos sido capaces de mejorar esta obra.
Colaboradores
COLABORADORES
David D. Eisenstat MD, MA, FRCPC, Professor and Chair,
Department of Oncology, University of Alberta, Muriel & Ada Hole
Kids with Cancer Society Chair in Pediatric Oncology; Professor,
Departments of Medical Genetics and Pediatrics, Faculty of
Medicine, University of Alberta, Edmonton, Canada
Jeffrey T. Wigle PhD, Principal Investigator, Institute of
Cardiovascular Sciences, St. Boniface Hospital Research Centre;
Associate Professor, Department of Biochemistry and Medical
Genetics, University of Manitoba, Winnipeg, Manitoba, Canada

REVISORES CLÍNICOS
Albert E. Chudley MD, FRCPC, FCCMG, Professor Emeritus of
Pediatrics and Child Health, and Biochemistry and Medical
Genetics, Max Rady College of Medicine, Faculty of Health Sciences,
University of Manitoba, Winnipeg, Manitoba, Canada
Michael Narvey MD, FRCPC, FAAP, Section Head, Neonatal
Medicine, Health Sciences Centre and St. Boniface Hospital;
Assistant Professor of Pediatrics and Child Health, Max Rady
College of Medicine, Faculty of Health Sciences, University of
Manitoba, Winnipeg, Manitoba, Canada

FIGURAS E IMÁGENES (FUENTES)

Agradecemos a los colegas que enumeramos a continuación las

imágenes clínicas que nos han prestado para este libro, así como su
autorización para usar figuras de sus trabajos publicados:
Steve Ahing DDS, Division of Oral Diagnosis and Radiology,
Faculty of Dentistry, Department of Pathology, University of
Manitoba, Winnipeg, Manitoba, Canada
Figura 18.10B-D
Vassilios Fanos MD, Department of Pediatrics, University of
Verona, Verona, Italy
Figura 19.3
Volker Becker MD †, Pathologisches Institut der Universität,
Erlangen, Germany
Figuras 8.12 y 8.14
S. Robben MD, Maastricht University Medical Centre, Maastricht,
The Netherlands
Figura 11.6B
David Bolender MD, Department of Cell Biology, Neurobiology,
and Anatomy,
Medical College of Wisconsin, Milwaukee, Wisconsin, USA
Figura 15.13A
Peter C. Brugger MD, PhD, Associate Professor/Privat Dozent,
Center for Anatomy
and Cell Biology, Medical University of Vienna, Vienna, Austria
Imagen de cubierta (resonancia magnética de un feto de nalgas)
Jack C.Y. Cheng MD, Department of Orthopaedics and
Traumatology, The Chinese University of Hong Kong, Hong Kong,
China
Figura 15.18
Albert E. Chudley MD, FRCPC, FCCMG, Department of
Pediatrics and Child Health, Section of
Genetics and Metabolism, Children’s Hospital,
University of Manitoba, Winnipeg, Manitoba, Canada
Figuras 5.12, 10.30, 12.17AB, 12.24, 13.13, 13.26, 15.24, 15.25, 15.26,
16.10, 16.11, 16.23, 17.14, 19.4, 19.5, 19.6, 19.9, 19.10, 19.12 y 19.14A
Blaine M. Cleghorn DMD, MSc, Faculty of Dentistry, Dalhousie
University, Halifax, Nova Scotia, Canada
Figura 18.10A
Heather Dean MD, FRCPC, Department of Pediatrics and Child
Health, University of Manitoba, Winnipeg, Manitoba, Canada
Figuras 13.17, 13.25 y 19.13
Marc Del Bigio MD, PhD, FRCPC, Department of Pathology
(Neuropathology), University of Manitoba, Winnipeg, Manitoba,
Canada
Figuras 15.10, 16.22 y 16.26
João Carlos Fernandes Rodrigues MD, Servico de Dermatologia,
Hospital de Desterro, Lisbon, Portugal
Figura 18.3
Frank Gaillard MB, BS, MMed, Department of Radiology, Royal
Melbourne Hospital,
Melbourne, Victoria, Australia; Radiopaedia.org
Figuras 9.8C y 10.17
Gary Geddes MD, Lake Oswego, Oregon, USA
Figura 15.13B
Bruno L. Vendittelli MD, New York University Medical Center,
Institute of
Reconstructive Plastic Surgery, New York, New York, USA
Figura 10.31
Christopher R. Harman MD, FRCSC, FACOG, Department of
Obstetrics, Gynecology, and Reproductive
Sciences, Women’s Hospital and University of Maryland,
Baltimore, Maryland, USA
Figura 12.16
Jean Hay MSc †, University of Manitoba, Winnipeg, Manitoba,
Canada
Figuras 7.2 y 7.4
Health Sciences Centre
Children’s Hospital and University of Manitoba, Winnipeg,
Manitoba, Canada
Figuras 10.13 y 19.7
Lyndon M. Hill MD, Magee-Women’s Hospital, Pittsburgh,
Pennsylvania, USA
Figura 12.5
Klaus V. Hinrichsen MD †, Medizinische Fakultät, Institut für
Anatomie, Ruhr-Universität Bochum, Bochum, Germany
Figuras 10.2 y 10.25
Evelyn Jain MD, FCFP, Breastfeeding Clinic, Calgary, Alberta,
Canada
Figura 10.22
John A. Jane, Sr. MD, Department of Neurological Surgery,
University of
Virginia Health System, Charlottesville, Virginia, USA
Figura 15.11AB
Dagmar K. Kalousek MD, Department of Pathology, University
of British Columbia;
Children’s Hospital, Vancouver, British Columbia, Canada
Figuras 12.12A y 13.10
James Koenig MD, FRCPC, Department of Radiology, Health
Sciences Centre,
Winnipeg, Manitoba, Canada
Figura 14.28D
Wesley Lee MD, Department of Obstetrics and Gynecology,
Division of
Fetal Imaging, William Beaumont Hospital, Royal Oak, Michigan,
USA
Figura 16.12A
Deborah Levine MD, FACR, Department of Radiology, Obstetric
and Gynecologic
Ultrasound, Beth Israel Deaconess Medical Center,
Boston, Massachusetts, USA
Figuras 7.5B, 16.12B e imagen de cubierta (resonancia magnética de un feto
de 27 semanas)
Mina Leyder MD, Universitair Ziekenhaus Brussels, Brussels,
Belgium
Figura 14.19
E.A. (Ted) Lyons OC, MD, FRCPC, FACR, Departments of
Radiology, Obstetrics & Gynecology, and
Human Anatomy & Cell Science, Division of
Ultrasound, Health Sciences Centre, University of
Manitoba, Winnipeg, Manitoba, Canada
Figuras 4.6B, 5.1, 5.10, 6.6, 7.1, 7.9, 8.4, 12.17CD e imagen de cubierta
(ecografía de un feto de 9 semanas)
Maulik S. Patel MD, Consultant Pathologist, Surat, India;
Radiopaedia.org
Figura 5.13
Martin H. Reed MD, FRCPC, Department of Radiology,
University of Manitoba and, Children’s Hospital, Winnipeg,
Manitoba, Canada
Figura 12.23
Gregory J. Reid MD, FRCSC, Department of Obstetrics,
Gynecology, and Reproductive
Sciences, University of Manitoba, Women’s Hospital,
Winnipeg, Manitoba, Canada
Figura 14.9
Michael and Michele Rice
Figura 7.6
Prem S. Sahni MD, Formerly of the Department of Radiology,
Children’s
Hospital, Winnipeg, Manitoba, Canada
Figura 15.14
Gerald S. Smyser MD, Formerly of the Altru Health System,
Grand Forks, North Dakota, USA
Figuras 10.17, 15.11C y 17.13
Pierre Soucy MD, FRCSC, Division of Pediatric Surgery,
Children’s Hospital of Eastern Ontario, Ottawa, Ontario, Canada
Figuras 10.10 y 10.11
Alexandra Stanislavsky MD, Department of Radiology, Mercy
Hospital for Women,
Royal Melbourne Hospital, Melbourne, Victoria,
Australia; Radiopaedia.com
Figura 12.12B
W. Jerry Oakes MD, Department of Pediatric Neurosurgery,
Children’s, Hospital, Birmingham, Alabama, USA
Figura 16.24
Edward O. Uthman MD, Consultant Pathologist,
Houston/Richmond, Texas, USA
Figura 5.3C
Elspeth H. Whitby BSc, MB, ChB (Hons), FFDRCSI, Academic
Unit of Reproductive and Developmental
Medicine, Department of Academic Pathology,
University of Sheffield, Sheffield, England, United Kingdom
Figura 16.25
Nathan E. Wiseman MD, FRCSC, Department of Surgery, Section
of Pediatric and
Cardiothoracic Surgery, Children’s Hospital, University
of Manitoba, Winnipeg, Manitoba, Canada
Figuras 9.8B y 12.15
† Fallecido.
Prefacio
Hace más de 46 años que publicamos Desarrollo embrionario. Esta
concisa obra está basada en nuestra otra obra, Embriología clínica,
11.ª edición, más extensa.
La 10.ª edición de Desarrollo embrionario se ha actualizado por
completo para reflejar los conocimientos existentes actualmente en
embriología clínica humana. Proporciona los fundamentos del
desarrollo normal y patológico. Como en ediciones anteriores, la
información clínica aparece destacada en cuadros de color verde. Cada
capítulo ha sido revisado meticulosamente con el fin de reflejar los
nuevos avances científicos y su relevancia clínica, así como nuevos
conocimientos en biología del desarrollo.
Hemos incorporado nuevas y numerosas fotografías en color de
embriones, fetos (normales y patológicos), neonatos (recién nacidos)
y niños. También aparecen muchas imágenes diagnósticas nuevas:
ecografías (sonogramas), TC (tomografías computarizadas) e IRM
(imágenes de resonancia magnética) de embriones y fetos.
Un rasgo importante de este libro son las Preguntas orientadas
clínicamente que aparecen al final de cada capítulo. Se incluyen,
asimismo, muchos estudios de casos clínicos de interés práctico,
disponibles en studentconsult.com*, junto con preguntas, respuestas
y explicaciones. Este material será de utilidad para todos aquellos
alumnos que quieran autoevaluarse y preparar exámenes.
Un conjunto innovador de 18 animaciones en color acompaña
esta 10.ª edición de Desarrollo embrionario, cuyo fin es apoyar a los
estudiantes en el aprendizaje de las complejidades del desarrollo
embrionario. Estas animaciones están accesibles en
studentconsult.com. Los profesores disponen también de
animaciones en alta resolución para su uso en las clases si han
adquirido o bien este libro o bien Embriología clínica (consulte al
representante de Elsevier). Cuando una animación presenta especial

interés para el texto del libro aparece este icono al margen.

La sección de teratología (estudio de las anomalías congénitas) ha
sido actualizada, pues es necesario el conocimiento del desarrollo
patológico para comprender las causas de las anomalías congénitas y
el modo en que pueden prevenirse. Se resaltan los aspectos
moleculares de la biología del desarrollo a lo largo de todo el libro,
especialmente en aquellas áreas que parecen prometedoras para la
medicina clínica y futuras investigaciones. Asimismo, el capítulo 20
se dedica en exclusiva a aportar información más detallada sobre las
bases celulares y moleculares del desarrollo.
Keith L. Moore
T.V.N. (Vid) Persaud
Mark G. Torchia

* Todos los contenidos de studentconsult.com están en inglés

Agradecimientos
Muchos colegas y alumnos han aportado valiosas contribuciones a
esta 10.ª edición del libro Desarrollo embrionario. Quedamos en deuda
con los siguientes colegas (nombrados alfabéticamente), bien por sus
revisiones críticas de los capítulos, por sugerir mejoras en el libro o
por proporcionar algunas de las figuras nuevas: Dr. Steve Ahing,
Faculty of Dentistry, University of Manitoba, Winnipeg, Manitoba;
Dr. David Bolender, Department of Cell Biology, Neurobiology, and
Anatomy, Medical College of Wisconsin, Milwaukee, Wisconsin;
Professor Margaret Buckingham, Department of Developmental
Biology, Pasteur Institute, Paris, France; Dr. Albert Chudley,
Departments of Pediatrics and Child Health and Biochemistry and
Medical Genetics, University of Manitoba, Winnipeg, Manitoba; Dr.
Blaine M. Cleghorn, Faculty of Dentistry, Dalhousie University,
Halifax, Nova Scotia; Dr. Frank Gaillard, Radiopaedia.org, Royal
Melbourne Hospital, Melbourne, Victoria; Dr. David F. Gomez-Gil,
Chicago, Illinois; Dr. Boris Kablar, Department of Anatomy and
Neurobiology, Dalhousie University, Halifax, Nova Scotia; Dr.
Deborah Levine, Beth Israel Deaconess Medical Center, Boston,
Massachusetts; Dr. Marios Loukas, St. George’s University, True
Blue, Grenada; Professor Bernard J. Moxham, Cardiff School of
Biosciences, Cardiff University, Cardiff, Wales; Dr. Michael Narvey,
Department of Pediatrics and Child Health, University of Manitoba,
Winnipeg, Manitoba; Dr. Drew Noden, Department of Biomedical
Sciences, Cornell University, College of Veterinary Medicine, Ithaca,
New York; Dr. Shannon E. Perry, San Francisco State University, San
Francisco, California; Dr. Gregory J. Reid, Department of Obstetrics,
Gynecology, and Reproductive Sciences, University of Manitoba,
Winnipeg, Manitoba; Professor Sir Peter W.J. Rigby, The Babraham
Institute, Cambridge, England; Dr. L. Ross, Department of
Neurobiology and Anatomy, University of Texas Medical School at
Houston, Houston, Texas; Dr. Michael A. Rudnicki, Regenerative
Medicine Program, Ottawa Hospital Research Institute, Ottawa,
Ontario; Dr. J. Elliott Scott, Departments of Oral Biology and Human
Anatomy & Cell Science, University of Manitoba, Winnipeg,
Manitoba; Dr. Gerald S. Smyser, anteriormente en Altru Health
System, Grand Forks, North Dakota; Dr. Alexandra Stanislavsky,
Department of Radiology, Mercy Hospital for Women, Royal
Melbourne Hospital, Melbourne, Victoria; Dr. Richard Shane Tubbs,
Children’s Hospital, Birmingham, Alabama; Dr. Edward O. Uthman,
Consultant Pathologist, Richmond, Texas, y Dr. Michael Wiley,
Department of Surgery, Division of Anatomy, Faculty of Medicine,
University of Toronto, Toronto, Ontario.
Los dibujos fueron realizados por Hans Neuhart, presidente del
Electronic Illustrators Group en Fountain Hills, Arizona. De la
editorial Elsevier, nuestro agradecimiento a Mr. Jeremy Bowes,
Content Strategist, por su valiosa percepción e intuición para dar
apoyo en la preparación de la 10.ª edición de este libro.
Agradecemos también a Ms. Sharon Nash, Content Development
Specialist, por su asesoramiento y sus muchas y útiles sugerencias.
Finalmente, agradecemos al equipo de producción de Elsevier,
especialmente a Mrs. Julie Taylor, Project Manager, por llevar a
término este libro. Esta nueva edición de Desarrollo embrionario es el
resultado de su dedicación y pericia técnica.
Keith L. Moore
T.V.N. (Vid) Persaud
Mark G. Torchia
Saca el mayor partido de la 10.ª
edición de Desarrollo
embrionario.

Con la compra de este libro tendrás acceso a una gran variedad de

MATERIAL COMPLEMENTARIO que refuerzan los contenidos del

libro impreso y te ayudarán en tu aprendizaje*. Este icono indica

qué secciones del libro van acompañadas de material electrónico
directamente relacionado:

• 18 magníficas animaciones, locutadas por expertos, para

guiarte a través de conceptos embriológicos clave:
Título de la animación Capítulo(s) asociado(s)
Fecundación 2, 3
Blastocisto 3, 4
Implantación 4
Gastrulación 5
Plegamiento del embrión 6
Cavidades corporales 9, 14
Aparato faríngeo 10
Cara y paladar 10
Aparato respiratorio 11
Tracto gastrointestinal 12
Aparato urinario 12, 13
Aparato reproductivo 13
Corazón 14
Sistema vascular 14
Desarrollo de los miembros 15
Sistema nervioso 5, 16
Desarrollo de los ojos 17
Desarrollo de los oídos 17

• Casi 700 preguntas de respuesta múltiple, con respuestas

razonadas, para ayudarte a comprobar tu conocimiento y a
preparar los exámenes
• Casos clínicos interactivos que acompañan a cada capítulo,
para ayudarte a relacionar el desarrollo humano y la práctica
 clínica diaria

No te pierdas todo este contenido adicional –mira en el interior

de la cubierta para obtener las instrucciones de acceso.
* Todos los contenidos de studentconsult.com están en inglés
1: Introducción al desarrollo
humano
El desarrollo humano comienza en el momento de la fecundación,
cuando un ovocito de una mujer es fecundado por un
espermatozoide de un hombre y se convierte en una célula única
llamada cigoto. El desarrollo comprende muchos cambios que
transforman el cigoto en un ser humano multicelular. La embriología
estudia el origen y el desarrollo del ser humano, desde el cigoto
hasta el nacimiento. Las etapas del desarrollo antes del nacimiento se
muestran en la figura 1.1.
 FIG. 1.1Fases tempranas del desarrollo humano.
Se muestran un folículo ovárico conteniendo un
ovocito, la ovulación y las fases del ciclo
menstrual.

Importancia y avances en embriología

El estudio de las etapas prenatales y de los mecanismos del
desarrollo humano nos ayuda a entender las relaciones normales de
las estructuras del cuerpo del adulto y las causas de los defectos
congénitos (anomalías congénitas). La práctica moderna de la
obstetricia se relaciona en gran medida con la embriología clínica o
embriología aplicada. Debido a que algunos niños presentan
anomalías congénitas, tales como espina bífida o enfermedades
cardíacas congénitas, los pediatras perciben fácilmente la
importancia de la embriología. Avances llevados a cabo en cirugía,
especialmente en los procedimientos relacionados con las etapas
prenatal y pediátrica, han hecho más relevante el conocimiento del
desarrollo humano desde el punto de vista clínico. Además,
paralelamente al descubrimiento de nueva información en relación
con los procesos del desarrollo, vamos comprendiendo mejor
muchas enfermedades, sus procesos y sus tratamientos.
Los nuevos progresos en biología molecular en laboratorios de
investigación han llevado al uso de técnicas sofisticadas (p. ej., la
tecnología genómica, los modelos quimera y transgénicos y la
manipulación de células madre) con el fin de explorar temas tan
diversos como la regulación genética de la morfogénesis, la
expresión regional y temporal de genes específicos, así como los
mecanismos por los que las células se especifican o se diferencian
para formar las diferentes partes del embrión. Los investigadores
continúan aprendiendo cómo, cuándo y dónde determinados genes
son activados y expresados en el embrión durante el desarrollo
normal y patológico.
El desarrollo comienza con la fecundación (v. fig. 1.1, primera
semana). El periodo embrionario comprende las primeras 8
semanas de desarrollo del embrión. El periodo fetal comienza en la
novena semana. El examen del calendario muestra que los progresos
visibles más externamente tienen lugar entre la tercera y la octava
semanas.
Se está conociendo muy rápidamente el papel crítico de genes,
moléculas señalizadoras, receptores y otros factores moleculares en
regular el desarrollo temprano del embrión. En 1995, Edward B.
Lewis, Christiane Nüsslein-Volhard y Eric F. Wieschaus recibieron el
Premio Nobel de Fisiología o Medicina por sus descubrimientos de
genes que controlan el desarrollo embrionario. Estos
descubrimientos están contribuyendo a una mejor comprensión de
las causas de los abortos espontáneos y las anomalías congénitas.
Robert G. Edwards (1925-2013) y Patrick Steptoe (1913-1988)
lideraron uno de los progresos más revolucionarios en la historia de
la reproducción humana: la técnica de la fecundación in vitro. Sus
estudios dieron como resultado el nacimiento de Louise Brown, la
primera «niña probeta», en 1978. Edwards fue galardonado con el
Premio Nobel en 2010.
En 1997, Ian Wilmut y sus colaboradores fueron los primeros en
crear un mamífero (una oveja llamada Dolly) mediante clonación,
usando la técnica de transferencia nuclear de células somáticas.
Otros animales han sido clonados con éxito desde entonces a partir
de cultivos de células adultas diferenciadas. El interés en la
clonación humana ha generado considerable debate debido a las
implicaciones sociales, éticas y legales. Además, hay preocupación
porque esta clonación pueda conducir a un aumento en el número
de recién nacidos con defectos del desarrollo y enfermedades
importantes.
 Las células madre embrionarias humanas son pluripotenciales y
capaces de desarrollarse en diversos tipos celulares. El aislamiento y
cultivo de células madre embrionarias humanas y otras células
madre pueden constituir una gran promesa para el desarrollo de
terapias moleculares.

Términos descriptivos
En anatomía y embriología se usan términos específicos de posición,
dirección y diferentes planos del cuerpo. Las descripciones en el
adulto se basan en la posición anatómica; el cuerpo está erguido, los
miembros superiores se encuentran a ambos lados y las palmas de
las manos miran hacia delante (fig. 1.2A). Los términos descriptivos
de posición, dirección y planos usados para el embrión se muestran
en la figura 1.2B a E. Cuando se describe el desarrollo es necesario
usar palabras que denoten la posición de una parte en relación con
otra o con el cuerpo en su conjunto. Por ejemplo, la columna
vertebral se desarrolla en la parte dorsal del embrión, mientras que
el esternón lo hace en su parte ventral.
FIG. 1.2 Esquemas de los términos descriptivos
de posición, dirección y planos del cuerpo. A,
Visión lateral de un adulto en posición anatómica.
B, Visión lateral de un embrión de 5 semanas. C
y D, Visión ventral de un embrión de 6 semanas.
El plano medio es un plano vertical imaginario de
sección que pasa longitudinalmente a través del
cuerpo, dividiéndolo en dos mitades, derecha e
 izquierda. Un plano sagital es cualquier plano
paralelo al plano medio. Un plano horizontal es
cualquier plano que forma un ángulo recto con
los planos medio y frontal. E, Visión lateral de un
embrión de 7 semanas. Un plano frontal (coronal)
es cualquier plano vertical que corta el plano
medio en ángulo recto y divide el cuerpo en
anterior (o ventral) y posterior (o dorsal).

Preguntas orientadas clínicamente

1. ¿Por qué estudiamos la embriología humana? ¿Tiene algún

valor práctico en medicina y otras ciencias de la salud?
2. Los médicos fechan un embarazo desde el primer día del
último ciclo menstrual, pero el embrión no comienza su
formación hasta unas 2 semanas después (v. fig. 1.1). ¿Por
qué usan los médicos este método?

Las respuestas a estas preguntas se encuentran al final de este

libro.
2: Reproducción humana
La pubertad comienza cuando aparecen los caracteres sexuales
secundarios, generalmente entre los 10 y los 13 años en las mujeres y
los 12 a 14 en los hombres. La menarquia (primer periodo
menstrual) puede ocurrir a una edad tan temprana como los 8 años.
En las mujeres, la pubertad ha debido completarse a los 16 años. En los
varones, la pubertad termina a la misma edad; finaliza cuando se
forman los primeros espermatozoides maduros.

Órganos reproductores
Los órganos reproductores producen y transportan células
germinales (gametos) desde las gónadas (testículos u ovarios) al
lugar de la fecundación en la trompa uterina (fig. 2.1).
 Secciones sagitales esquemáticas de las
FIG. 2.1
regiones pélvicas de una mujer (A) y un hombre
 (B).

Órganos reproductores femeninos

Vagina
La vagina sirve para el paso del flujo menstrual, aloja el pene
durante el coito y forma la parte inferior del canal del parto,
constituido por la parte del útero y la vagina a través de los cuales
sale el feto (v. fig. 2.1A y B).

Útero
El útero es un órgano de pared gruesa con forma de pera (fig. 2.2A y
B) que consta de dos partes principales:

• El cuerpo uterino, que constituye los dos tercios superiores.

• El cuello uterino, formado por el tercio inferior.
Another random document with
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Street would either drift into his arms or have ‘to step down and
out’—to abdicate the crown she has worn so long.” Vreeland
lumbered along, building up fanciful solutions of the mystery.
In the now almost incessant “duty service” near his beautiful fiancée,
Vreeland a hundred times endeavored to trace back James
Garston’s early life. But the blue-eyed Nixie who was soon to be his
wife only laughed merrily.
“Pray remember, sir, that Senator Garston is my guardian. After my
dear father’s death, my mother went abroad, and I was educated in
the ‘Sacré Cœur’ Convent at Brussels. Her death left me alone in the
world.
“‘Uncle James’ had been almost forgotten by me in the thirteen years
which we passed in Paris and Brussels, and as I left the West a
mere child, all my memories are the vanishing dreams of childhood.
All his social past is a sealed book to me.”
Vreeland was fain to be content, as the lovely ingenue concluded:
“All I know is that he has always managed my affairs, and that his
personal history is linked with the development of the whole region
west of the Rockies. Why, you should know his history from your
own Western wanderings.”
“Was he ever married?” timidly hazarded Vreeland. But, the young
society queen only laughed back.
“Ask him! And then ponder now the possibility of another marriage.
You are now, sir, to take me driving. The only marriage which
concerns you, is a joint affair.”
That afternoon, as they drove through the park under the
chaperonage of the amiable Mrs. Volney McMorris, Vreeland
unsuccessfully endeavored to allay his recent dissatisfaction at the
absence of any womanly background for the highly polished
“Western diamond,” which he was soon to win and wear for life.
The story of the young heiress was smooth enough and faultlessly
delivered. Vreeland forebore to “pump” Mrs. McMorris, for he was
well aware that she was “all things to all men,” and her voluble
explanations would carry no real conviction.
“She helped Alida Hathorn on to the very verge of ruin,” he gloomily
recalled.
“There might have been a marriage between myself and Elaine but
for her vicious intermeddling.
“She took that Isle of Wight story in commission and spread it all
over New York, while working both sides for coin—a woman Judas!”
While he returned the salutations of Messrs. Merriman, Wiltshire and
Rutherstone on the social parade, he was vaguely reflecting on the
uselessness of his crime as regarded the stealing of the hidden
paper and the tapping of the private wires, as well as the mail frauds.
It now followed him like his own shadow, and the paper was a source
of countless nightmares. If it were only safe!
“All that is useless now,” he growled. And he suddenly saw that he
was left in the power of Doctor Hugo Alberg, of Justine and of
August Helms, the janitor.
“There will be no speculation in ‘Sugar’ for months; the market is
dead, pending the reorganization and New Jersey reincorporation.
“My strange employer is away. She will not be here for months; and
she has also taken alarm at the presence of Garston.
“The whole lot of them will probably operate in a blind pool now.
There will be nothing for me to gain, and everything to lose in
running any further risks.”
He saw with concern that Alberg greatly missed his wealthy and
generous patient, and a few significant hints had proved to him that
the German physician was now “money hungry.”
“There is Justine always to be pacified, and that brute, Helms, too;
he will surely want money.
“Once married, and a fixture here, I am ‘nailed to the cross’ for
torture by these people—if they should turn against me.
“Fear will control Doctor Alberg at the last,” reflected Vreeland. “He
has been guilty of half-poisoning his patient.
“Justine I can surely rely on as long as I keep her pacified, but, that
brute Helms is steadily increasing in his money demands. Some
night, when drunk, he may blow the whole thing abroad.” And he had
caught a glimpse of Helms and Bagley diving into a saloon together.
It frightened him.
It was true that Helms had found his way down several times to the
Elmleaf to get money, in a half-fawning and half-threatening bluster.
And on several occasions when Vreeland was absent, the grave-
faced valet, Bagley, had joined the janitor, and in some hours spent
over the cups of Gambrinus had gained pointers which had given the
lively roundsman, Dan Daly, some very valuable hints.
There was in his cup of “bittersweet,” however, one great consolation
to the successful Harold Vreeland, whom all men now envied.
The impending union with Katharine Norreys would found his
fortunes on a solid basis; he would have the absolute protection of
the great speculative Senator, and the reports of his detectives told
him that Hugh Conyers was simply buried in his journalistic duties. It
seemed to be a lull in the war, even the pickets had ceased firing.
There were no conferences with Judge Hiram Endicott, and nothing
to indicate any activity among Romaine Garland’s friends.
Only one side of the whole affair remained dark to Vreeland. Even
Justine Duprez could not tell him how or why Elaine Willoughby had
openly taken her unacknowledged daughter to her house for shelter.
It was as yet a mystery as to whether fear, intrigue or accident had
brought the lovely girl into the opened arms of her still beautiful
mother.
“All I know,” said Justine, in a conference arranged for this purpose
by her now indifferent fellow-conspirator, “all I could find out was,
that this green-eyed cripple, this little sycophant Irelandaise, who
now is my tyrant, brought the tall girl late one evening to the
‘Circassia.’”
“It was a strange visit,” murmured Justine, “for she brought no
luggage, and that girl never left my mistress’ presence for a moment,
till she went away with the two Conyers.
“I am certain that Madame had never seen this girl in the seven
years of my employ. There were no pictures, no relics of childhood—
nothing. And I was always on the lookout for the mystery of
Madame’s life—”
Justine demurely dropped her eyes.
“Bah!” she cried; “a woman with blood as cold as a fish! No life, no
love; she cares for nothing but money.
“Among all of them, not a lover! I thought she was fond of the dead
Mr. Hathorn once, but he was soon on a level with the others.”
Justine’s voice was duly scornful.
“And then her tears and frequent fits of sorrow! That was the record
the whole of seven years.
“The last thing I saw of her—a stolen glance—she had this girl’s
picture in her hand, and was weeping over it.
“If she is a child of hers, she is probably a child of shame. She now
fears the exposure, and has gone abroad to hide the girl away
forever. Trust to Justine’s experience! I know these women saints.
They always have nibbled at le fruit defendu—hypocrites!”
Mr. Harold Vreeland fancied that he saw light at last. “I believe that I
can observe Senator Garston’s game. He would use this hidden fact
to force Elaine Willoughby into his arms. By Jove! she does fear him!
Perhaps Justine is right.
“And so, when I am married to Katharine, and Garston is free of all
social claims, if he alone knows her secret, it may be buried forever
in her marriage with him.
“To bring the proper pressure to bear, he must have the girl first. And
he would not be too good to bribe the girl with a fancied inheritance.
Once that the child is under his influence, Elaine’s proud heart must
either bend or break.
“For he will win his way to her side, even across the fires of Alynton’s
hate or the social ruin of Elaine’s good name.” Vreeland already
knew the iron will of the man who was driving ahead with
recklessness in the chase.
And so, armed with the deadly secret of the enormously powerful
cabal, the stolen document, Vreeland now knew that if brought to
bay, Elaine would perhaps be sacrificed by the secret syndicate,
despised by the undeceived Alynton, and then, with the secret of her
early life in Garston’s possession, be utterly at his mercy. “Yes, she
is in the toils,” he muttered. “There is no escape for her.”
It was at the wish of Senator James Garston, now lavishly liberal in
his preparations for his ward’s wedding, that the bridal was
postponed to the first days of June.
“All is going on well, Harold,” said Garston. “We have worked into a
thorough accord with all her representatives.
“And you will not find love-making with Katharine Norreys an irksome
task. I wish only to wait till I learn that Elaine Willoughby has landed
at Brindisi.
“Somewhere on the Continent she will surely meet this girl. I shall
have instant reports from my detectives. For so far, we have found
out Elaine’s route, but, the girl is still hidden.
“I wish you to go away at once on your wedding tour, and then to
keep Mrs. Willoughby in sight—within touch. I only want to meet the
mother and daughter face to face—only once. I will have my innings
then, and finish the whole matter in short order.” His face was
merciless now.
“Now, you will be no object of suspicion on your wedding tour; such a
happy voyage always explains itself,” he sardonically smiled. “The
moment that I am cabled for, I shall depart incognito. My work will be
quickly done when I find this sly woman and her child together. The
whole world is not wide enough to hide that child from me.” And
Vreeland drifted daily under Garston’s strong control; he was floating
with the tide, drunken with all his successes.
The days drifted along in all the preoccupation of daily business and
the growing bustle of the impending wedding.
Harold Vreeland was most agreeably surprised in the later days of
May by a cordial letter from Mrs. Willoughby, posted at Port Said.
Her congratulations upon his impending marriage were coupled with
her carte blanche as to leave of absence from the firm, and the
significant direction to leave Bagley in charge at the Elmleaf.
“We shall have business uses for the apartment during the winter,
and Miss Kelly will give Bagley all his orders and attend to the
accounts. I have directed Judge Endicott to present in my name to
your wife a proper reminder of the esteem which I have for her.”
The notification three days before the wedding, through Noel
Endicott, that Mrs. Willoughby had placed a year’s salary at his
personal disposal on the books of the firm, as an extra bonus,
carried away the last vestige of Vreeland’s haunting fears.
Nothing remained of the awkward episode of the inquiry as to the
stolen document, and Vreeland had already settled with Doctor
Alberg, and Helms with an affected liberality, for his absence.
Now socially entirely in the hands of Messrs. Wiltshire, Merriman and
Rutherstone, his three groomsmen, and having seen the resplendent
Mrs. Volney McMorris rally many beautiful Ishmaelites, married and
single, around his bride, Vreeland was moved forward to the altar on
the golden flood of Senator Garston’s splendidly liberal preliminary
entertaining.
The Western millionaire was touching up every cloud hanging over
Katharine VanDyke Norreys’ social haziness with a golden lining.
There remained but two things for the happy groom to do now.
The one was to have a last interview with Justine, who was now
reduced to a calm subserviency to the orders of the young “Private
Secretary,” and the other to effect a safe deposit in some satisfactory
place of the stolen document and its tell-tale copy.
He had decided to be liberal with Justine in money matters, and to
entrust her in his three months’ absence with the watching of Helms,
the janitor, and the disgruntled German doctor.
A famous plan suggested itself! Justine should feed out to these men
money, in his name, during his absence.
“And that, with the hope of more, will keep them true to me, as
rascals go, till I return.” He had once decided to dismantle the secret
connections with Mrs. Willoughby’s telegraph and telephone. It was
the subject of a long, introspective reverie.
But reflection had told him of a possible mistake. And perhaps in his
absence, Justine might glean from the detained correspondence
delivered at the “Circassia,” some facts to guide both Senator
Garston and himself. Yes, the “underground railroad” should not be
disturbed. Its existence was as yet concealed from all his enemies.
The use in the next winter of the “Elmleaf” rooms for a concealed
headquarters of speculation caused him to leave the wires in
position. “It might excite these people’s suspicions. I must appear to
trust them,” he decided, “and Garston may even make a million over
the private tips I can give him if I am up to their game.”
Suddenly it occurred to him that his own marriage might change the
situation, and yet, there were Elaine Willoughby’s recent orders.
“She means probably to hide her child, and then come back and be
Queen of the Street again,” he smiled. “The ruling passion. She has
the speculative mania still.” For it was clear to him now that the
presence of mother and daughter together in New York City was an
unnecessary risk.
And so, even on the threshold of his marriage Harold Vreeland
feared to trust his bride with the secret of the stolen document. They
were to live at the Hotel Savoy on their return, “so as to be near
Uncle James, at the Plaza.”
With a moral cowardice which he could not explain, Vreeland had as
yet declined to face the burning question of the stolen document.
The copy he had always carried secreted within the waistcoat lining
of his traveling suit. “I can easily leave that over in Europe,” he
murmured. “The original. Where shall I hide it?” He was long in the
dark.
But it was by a devilish impulse, aided by accident, that he found a
place in Justine Duprez’s rooms on South Fifth Avenue to safely hide
the dangerous original.
One of the plates of a door framing had sprung partly loose. A
sudden idea seized him. Her rooms were the safest place for many
reasons.
To gain time for preparation, he sent the old hag away on an errand.
Sealed in a cloth envelope, the paper was soon hidden behind the
upper framing plate, and with a hammer, covered with his kid gloves,
he drove the half-dozen old, rusted nails tightly home. And he gazed
in triumph at the neat device.
“They will of course think that she stole it, should it ever be found,”
he mused triumphantly, as he lit a Henry Clay and gloated over his
cunning.
“If the house should burn I am safe. In every way it would go up in
flames. If I should die, then it makes no difference to me what
happens. If she is caught—this would be damning evidence only
against her.
“And I would never dare to trust myself with either Garston or my
wife, and be found out in the custody of that document.
“Accidents will happen; I might fall ill, and now no matter what
befalls, it never can be traced to me.”
He grinned with joy as he contemplated depositing the copy abroad,
under an assumed name.
“It will there be safe from all American legal process, and the original
is here where I can use it if needed, and as it is, it can never be
traced to me.”
He carefully examined the exterior of the row of solid brick
tenements. They were good for a life of fifty years.
As he walked away, when he had “finished his letters,” and left a last
greeting for Justine, he stood upon the heights of an impregnable
position.
“It was a stroke of genius, that last idea of mine!” he gaily cried, as
his eye rested on an old woman who had just descended the stair.
He knew not the burden of her eager soul. She carried his fate!
Once around the corner, that old woman scuttered away to find
roundsman Dan Daly, for the peep-hole had covered a keenly-
glittering eye, even after Justine had left her sighing lover to his “last
bachelor letters.” And thus the hiding-place was known to more than
one.
But Vreeland hastened away in a triumphant glow of satisfaction.
The splendors of the Grace Church wedding, the gilded festivity of
the Waldorf wedding dinner, and all the countless preoccupations of
the impending voyage busied Harold Vreeland’s excited mind for
three days.
There were hundreds of valuable wedding presents to deposit in
safety, for society had showered gifts upon the successful interloper
with its hard-hearted, hollow flattery of success. It had been a
“society event,” and his face, with that of the beautiful bride, had
ornamented several “up-to-date” journals.
The flower-decked bridal staterooms of the “Campania” had received
Vreeland’s party, and Messrs. Rutherstone, Merriman and Wiltshire
were joining the bride and bridesmaids in the parting “loving cup,”
the table was covered with journals filled with the usual “glowing
accounts” and piled up high with congratulatory letters and
telegrams, when “Uncle James” drew the complacent bridegroom
aside.
In a private nook, he turned a scowling face to the happy Vreeland.
A yellow telegraph envelope fluttered from his hand to the desk as
he read again these disquieting words:
“She has telegraphed for a cabin on the ‘Normandie,’ and is coming
home alone. Took a special train from Vienna to Havre. All traces of
girl lost.”
“Vreeland,” growled the maddened man, “some one has betrayed
us. Wait at the Hotel Cecil, London, for my cipher orders.
“That woman is a devil in artfulness, and it is a fight to the death
now.”
Ten minutes later, the “Campania” was plowing down the beautiful
bay.
 CHAPTER XIV.

FOR THE CHILD’S SAKE!

The crowding passengers lounging on the decks of the “Campania”

and “Normandie” idly watched the fleeting waves torn up by the
ocean racers as they swept by each other in mid-ocean four days
later, but there were strangely agitated hearts, too, on the passing
steamers, when the signal flags were broken out.
For, the secret enemies now swept past each other at the distance of
a few furlongs.
“What the devil can the real motive of her quick return be?” angrily
mused bridegroom Vreeland, as he called up again Senator
Garston’s baffled fury on learning that for all his goading on, his
detectives had failed to locate the missing Romaine Garland.
He led his beautiful bride back to her room, and then left her to the
enjoyment of “Les Denis-Vierges,” while he eyed the fast-receding
“Normandie.”
“Another big deal in ‘Sugar,’” he suddenly thought, and he felt
himself perhaps hoodwinked by both Senators and the handsome
woman who had so artfully led him on to his fate. “It may be that they
all are fooling me; I may have been merely jockeyed away.
Mrs. Willoughby can work the ‘off side’ of her deals alone from the
‘Elmleaf,’ and the regular transactions will go on as usual through
our firm, really Alynton & Willoughby. Or, she may have picked up
another protégé. God only knows what a woman may do.
“They all have their secrets, by Jove! Senator Garston or this cool
devil, Hugh Conyers, may now turn up as the secret broker in my
place.”
It suddenly occurred to him that the powerful Western millionaire
might really be the favored lover, and Alynton, after all, only the dupe
of a growing passion. “I am powerless to go further now,” he
groaned, as he gazed at the rooms where his lovely and exacting
bride was “squeezing the orange of life” to its last drop. He had
found out, even now, that there were thorns upon his rosebud.
He was not yet entirely satisfied with the status of husband so
recently assumed. Still affecting all the delicacy of the lover, he had,
however, quite practically approached the subject of Katharine
Norreys’ investments “in the hands of Uncle James.”
And he soon found out that the exquisite form of his dazzling blonde
wife hid a resolute and undaunted spirit, an unruffled temper, and an
easy, natural defiance of all marital control. “Where did she get her
experience of life?” mused the startled bridegroom.
“You must go over all these tiresome matters, Harold, with Uncle
James, on our return,” the overwearied, fashionable bride answered.
“I have never entered into any details with him, and I supposed, of
course, that you and he had covered all this ground. I have only
asked him for money as I needed it since my return, and he has
always sent me his checks. It is for you, both business men, to
regulate such matters.” And she cast her eyes down again on her
entrancing book.
“Then you have no permanent bank account of your own?” moodily
demanded Vreeland.
“Why should I have one?” innocently replied Mrs. Katharine
Vreeland, “when Uncle James has always paid the bills and
furnished me all that I ask? I have never asked him for any formal
accounting.” Harold Vreeland was secretly nettled at her easy
carelessness.
“And if he were to die, if anything happened, you would then know
nothing of your own affairs,” said the dissatisfied husband.
“No more than I know now of yours, my dear,” calmly answered
Katharine, settling herself deeper in her cushions. “Uncle James
simply told me that you were a very rich man, and of course, I took
his word. I have not asked you to inventory your own possessions.”
She was turning an unusually interesting leaf as Vreeland walked out
of the cabin in a suppressed rage.
“We are both at sea, it appears,” was his disquieting thought, and
again the remembrances of that slender family tree of his lovely wife
annoyed him. It seemed to begin and end in the graves of the dead
parents, who were only gruesome shadows.
“I will go over this whole ugly matter with Garston at once, just as
soon as I see him,” was Vreeland’s mental decision. “Katharine is
either a child-wife of the Dora order, or else far deeper than the sea
that we are skimming over now.”
It came to him cogently that he had taken her “on trust” largely, and
that a current of life’s mysterious undertow had swept him along into
Senator Garston’s power. There was no going back, however.
“It is too late to hesitate now,” he mused, as he uneasily gazed back
toward America, well knowing that some giant game might be played
in his absence.
In the deal there would be no cards for him, however the luck might
turn. And there remained but one golden gleam in the gray clouds.
He had that paper with which to dominate Mrs. Willoughby. But, it
was a dangerous weapon; it might prove a boomerang.
“Justine Duprez stands between me and all harm. That was a
master-stroke! And so I can cut into the game as I wish, on my
return. The very first thing I shall do will be to get Katharine’s fortune
out of Garston’s control. He shall face the music. And yet, I can
afford no quarrel until that is all safe.”
In the month which followed this vain attempt at probing the financial
resources of the wife of his bosom, Mr. Harold Vreeland, at the Hotel
Cecil, London, found the beautiful Katharine’s money-spending
power to be something abnormal.
There was a rapid exchange of letters and cable ciphers between
Garston and the young broker spy, but the husband was never
enlightened as to the nature of the frequent telegrams and letters
passing between “Uncle James” and his ward.
It vastly annoyed him—this continued private commerce of ideas.
The questions of the husband were frankly enough met. “I have
always been accustomed to do exactly as I pleased,” the lady
remarked, with a bright, hard smile. Vreeland’s face hardened.
“And now, that you are married?” demanded Vreeland, angrily.
“I shall continue to do so, Harold,” his wife sweetly replied.
“If you would have me lead a Darby and Joan life, please to
remember that sort of thing went out with the ‘Rollo books’ and ‘Faith
Gartney’s Girlhood!’”
Mr. Harold Vreeland, the husband of a few weeks, soon realized that
while he was doing the clubs and music halls of London, his
resplendent wife had quietly gathered up quite a coterie of admiring
American men, generally conversationally lumped as “the Western
gang.”
These ardent cavaliers seemed to be all wifeless, and, strangely
enough too, without mothers or sisters. “‘Uncle James’ friends,” was
Mrs. Vreeland’s saving clause, when at last her angered husband
remonstrated at their increasing circle. He was beginning to be
agnostic as to her guilelessness.
And on their removal to Paris, where certain of these “friends” soon
after appeared, Katharine Vreeland bravely continued “to do as she
pleased,” and her now bitter husband partook himself to sparkling
wine and “the sights of Paris.”
He was driven along from day to day, for he had no reliable news
from the seat of war. He realized that he was alone in the world and
without one trusty friend. His wife was only a bright enigma.
“The lone-hand game has its disadvantages, I perceive,” was his
bitter secret comment, as he tired of the Hotel Continental—the
perfunctory drives in the Bois, the open summer amusements—and
visibly fretted at his wife’s endless shopping.
Even with Garston’s substantial bribe, he began to see that
Mrs. Katharine Vreeland’s “separate estate” was to become a very
“burning question”—in the near future.
She was a “money-eater” of the first class.
“Let us get back to New York,” he moodily said after one of a series
of wordy recriminations. “With all my heart,” placidly retorted the
“beautiful Mrs. Vreeland,” for she had now acquired that professional
designation in the journals and the cant phrases of the uneasy
floating “American circle” of Parisian high life.
Harold Vreeland was now mentally tired of the by-play of marital
fencing. He realized, in all their varied encounters, that she was
calmly superior at every clash.
Bright, bold and ready, she “came back at him” every time, and he
was quietly cornered by that flashing rapier, her tongue. What man
can prevail against that two-edged sword?
But one resource was left. He had run the gamut of sullenness,
persuasion, a bit of bullying, some pleading and even a touch of lofty
tenderness, but her point was carried high, her wrist easy, and her
blade opposed to him at every turn.
He could not avouch himself a mere fortune-hunter, and so, he took
refuge in an ominous and expectant silence. “I will get hold of her
estate, and then curb her extravagance,” he brooded.
His worst fears as to the “underground railroad” communications of
the “uncle” and ward were realized when he finally received a
positive request of Senator Garston for an immediate return.
“I want you at once. I wish to lay out our plans for the winter. And if I
am to trap this underhanded, intrigant Mrs. Willoughby, I must finish
my work before the opening of the session of Congress, and our
committees will begin soon to meet. Come on, with no delay.” The
words were almost mandatory, and they annoyed him strangely.
Returning from his banker’s with this letter, he found his wife’s two
maids busied in packing up all her effects. He was startled, but took
the defensive.
Something impelled him to keep the news to himself. “I am tired of
Paris,” shortly said his wife, as she recognized the drifting odor of an
absinthe frappée. “We can just catch the Gascogne, and so, I have
ordered all my bills sent in. You must attend to them, and then,
secure our passage.”
“Let me know their probable amount,” gruffly answered the husband,
as he departed for the steamer office. He was beginning to feel a
master hand now.
“She had the news before I received it,” he growled. “And I swear I
will make it my pleasing duty to bring ‘Uncle James’ to book, on my
return. I will get her property into my hands, and control it.
“She would beggar even a Vanderbilt, an Astor or a Goelet, if given a
free hand.” Vreeland aspired to the conquest of this defiant beauty in
rebellion.
It so happened that the game as laid out by “Uncle James” suited all
three; but, while he thirsted to see Justine Duprez once more and to
confer with Doctor Alberg, Vreeland was really anxious at heart to re-
enter the comparative protection of his Wall Street office.
“By Jove! I am at least between the lines there,” he mused. “I can
frighten both sides, and so, guard myself.”
It was on the Gascogne that he watched Katharine VanDyke Norreys
as the Count de Millefleurs (a young attaché going over on his first
appointment) bent over her steamer chair.
“This marriage has only hung a millstone around my neck,” he
resentfully brooded. “And I wonder if I was only brought in to relieve
‘Uncle James.’” It was a mean suspicion, but it clung closely to him.
He was now the prey of ugly thoughts, and fleeting fears disturbed
“the sleep of Richard.”
There were times when he feared for the safety of the document so
deftly hidden away. The copy had been artfully deposited (under
receipt) in a Belgian branch bank in Paris, under an assumed name,
and the banker’s receipt was now sewed in his waistcoat. “Thank
God! That is all safe!” he sighed.
He little reflected that one day, laughing over the “Agony Column” of
the London Times, his eye had paused at the name “Martha Wilmot.”
Some trace of familiarity, some fleeting memory caused him to read
the few lines.
“Handsome reward and the most complete immunity guaranteed.
Greatly to your advantage. Communicate in any way.”
The signature, “New York,” followed by an address, closed the
expensively placed announcement.
“Some relic of man’s folly and woman’s frailty!” he laughed. “The old,
old game goes on forever.”
And yet, he little dreamed that Hugh Conyers and handsome Dan
Daly were now the right and left hand men of Judge Hiram Endicott,
who was engaged in some very interesting metropolitan researches.
In far-away New York, there was the veiled duel of two fearless
intellects going on, even in the summer days, when the town was
empty.
Mrs. Elaine Willoughby was again the radiant mistress of Lakemere,
although she spent a portion of her time in town at the Circassia.
There was now a strange glow of happiness shining on the splendid
woman’s face, and the services of Doctor Hugo Alberg were
permanently discontinued.
It was impossible for the revengeful Teuton to learn the reason from
Justine Duprez. The courteous terms of Mrs. Elaine Willoughby’s
letter, inclosing a check for his annual account, were too
unmistakable to be misconstrued even by the dense German. It was
a congé not to be misunderstood. His Waterloo!
And, in a roundabout way he had also learned that Judge Endicott
and his nephew made up the whole social circle at Lakemere, with
Hugh Conyers as a permanent summer guest.
Hugo Alberg had sworn an oath that Harold Vreeland should recoup
him for the loss of his star patient. He now only awaited the return of
his proposed victim “to levy the Rhine dues.”
A visit to the South Fifth Avenue rooms where Justine had
vicariously entertained him in the old days, gave him the news, by
the mouth of the old denizen, that “la pauvre Justine” was tied down
at Lakemere.
“Some one have robbed ze lady last year, and now Justine is ze
prissonaire to watch ze garderobe all ze while; and only ze travail
and ze solitude! V’la tout! Pauvre Justine! Elle vent bien partir pour la
France.” The doctor hungrily awaited Vreeland’s return for a bleeding
process.
No one but the Frenchwoman herself knew how tightly the coils were
wound around her. Shaking in fear, left without the secret protection
of her traitorous tempter, Vreeland, she dared not try to break away
from Lakemere, for she now feared the gleaming wrist-irons.
To run away would be only to invite an instant arrest, and she panted
for the time of the winter’s gaieties. She would have a chance
perhaps then to slip away unknown.
Her plan was already formulated. A simulated illness, a last
“bleeding” of Harold Vreeland, and then, a return to dear Paris. Once
again on French soil, she would be safe. For Paris would soon
swallow her up. The vicious child would be hidden in the mighty
bosom of the Mother of all Wickedness.
“Ah! he shall pay,” she muttered, as her velvety eyes rested, lit up
with a strange fire, on the beautiful woman whose iron hand now
held her so firmly. “She and the Kelly—how I could drive a knife into
their hearts!” she hissed.
“But Justine must wait; gold first, gold—and then la liberté shall be
mine.”
When “Harold Vreeland and wife” were duly domiciled at the Hotel
Savoy, he was not astonished at the proximity of “Uncle James” at
the Plaza Hotel; but, even on the pier, when the Senator met them,
Vreeland noted the ravages of some overmastering passion in the
strong man’s face.
The eyes were brilliant and unsteady, there was a foreign irritability
in his abrupt manner, and Vreeland’s attempts at a tête-à-tête were
only met with a sharp command “to get inside his old business lines”
as soon as he could; and Vreeland, humbled, kept his temper.
“I must have you back in the traces again,” sharply cried Garston.
“And, I would get up to Lakemere to-night if I were you. See
Mrs. Willoughby, and get safe on the old basis.
“The stock market is humming, and I will soon have need of you in
Wall Street. I trust no one there but you.”
Harold Vreeland hastened away to the office, and found the same
unimpassioned greeting which had always characterized Horton
Wyman. And in the rush, they were now glad to have his aid in their
increasing affairs.
“You will go, of course, up to Lakemere to-night?” said Noel Endicott.
“I have already telegraphed your arrival to Mrs. Willoughby.”
In a stolen detour, Vreeland arranged for an early morning interview
with Doctor Alberg, and then he passed the “Circassia” on his way to
the train after dinner.
The flat demand of janitor Helms for “backsheesh” keenly angered a
man already enraged by “Uncle James’” quiet appropriation of the
first evening with that hawk-eyed free-lance of marital beauty,
Mrs. Katharine Vreeland, “whose remarkable loveliness had created
such a London and Paris sensation.”
“I will soon cut the Gordian knot between these two,” growled
Vreeland, as he descended from the waiting carriage at Lakemere. “I
will either have my wife and her property to myself, or else ‘Uncle
James’ will show his hand, to the very last card.” He was beginning
to be reckless in a blind jealousy.
The welcome of Mrs. Elaine Willoughby to her returned protégé was
merely a complacently cordial one, and yet, in half an hour, Vreeland
bore away the assurance of lulled suspicions and his continued
business relation.
“I shall soon call upon Mrs. Vreeland and assure myself by
inspection of her married happiness,” was the last greeting of the
hostess, whose other guests, if any, were invisible.
“I will send for you to the ‘Circassia’ next week, and give you my
general directions for some business which is impending.”
“That woman has found a new happiness. Her life is now complete,”
was the keen-eyed schemer’s comment as he sauntered away
toward the park gates, where the impatient horses awaited his
return.
A flitting form in the dusky garden walks led him toward the “lovers’
labyrinth,” behind the unforgotten summer house. His one friend was
on watch.
“Justine!” he gasped, and he hastened to stealthily join her in the
deepened gloom of the trees. A new fear smote upon his startled
nerves.
There was the velvet-eyed Frenchwoman in waiting, and her
passionate words, her panting breast and gleaming eyes told him of
an unbroken tie, the bond of their guilty past.