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General Urology
NINETEENTH EDITION
Edited by
Jack W. McAninch, MD, FACS, FRCS(E)(Hon)

Professor of Urology
University of California School of Medicine
Chief, Department of Urology
San Francisco General Hospital
San Francisco, California
Tom F. Lue, MD, FACS, ScD (Hon)

Department of Urology
University of California School of Medicine
New York Chicago San Francisco Athens London Madrid Mexico City
Milan New Delhi Singapore Sydney Toronto
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Contents
Contributors vii 10 Laparoscopic Surgery 149
Preface xi
David B. Bayne, MD, MPH;
J. Stuart Wolf, Jr., MD, FACS;
1 Anatomy of the Genitourinary Tract 1
Marshall L. Stoller, MD; & Thomas Chi, MD
Emil A. Tanagho, MD; &
Tom F. Lue, MD, ScD (Hon), FACS 11 Robotic Surgery in Urology 167
Maxwell V. Meng, MD, MPH
2 Embryology of the Genitourinary System 17
Emil A. Tanagho, MD; Hiep T. Nguyen, MD; 12 Urinary Obstruction & Stasis 177
& Michael DiSandro, MD
Marshall L. Stoller, MD; &
3 Symptoms of Disorders of the
Genitourinary Tract 31
13 Vesicoureteral Reflux 191
Benjamin N. Breyer, MD, MAS, FACS
Thomas W. Gaither, MD, MAS; &
Hillary L. Copp, MD, MS
4 Physical Examination of the
Genitourinary Tract 41
14 Bacterial Infections of the
Maxwell V. Meng, MD, MPH; & Genitourinary Tract 201
Emil A. Tanagho, MD
Mary K. Wang, MD; &
Hillary L. Copp, MD, MS
5 Urologic Laboratory Examination 49
Anobel Y. Odisho, MD, MPH; 15 Specific Infections of the
Sima P. Porten, MD, MPH; & Genitourinary Tract 229
Kirsten L. Greene, MD, MS
Emil A. Tanagho, MD; &
Christopher J. Kane, MD, FACS
6 Radiology of the
Urinary Tract 63
16 Sexually Transmitted Infections 243
Daniela Franz, MD; Scott Gerst, MD; &
Hedvig Hricak, MD, PhD Kristin Madden, PharmD;
Amanda B. Reed-Maldonado, MD, FACS;
& John N. Krieger, MD
7 Vascular Interventional Radiology 107
Ryan Kohlbrenner, MD; & Roy L. Gordon, MD 17 Urinary Stone Disease 259
Marshall L. Stoller, MD
8 Retrograde Instrumentation of
the Urinary Tract 117
18 Injuries to the Genitourinary Tract 291
9 Percutaneous Endourology and
Ureterorenoscopy 129 19 Urothelial Carcinoma: Cancers of the
Bladder, Ureter, and Renal Pelvis 309
David B. Bayne, MD, MPH;
Joachim W. Thüroff, MD; Badrinath R. Konety, MD, MBA; &
Rolf Gillitzer, MD; & Thomas Chi, MD Peter R. Carroll, MD, MPH
iii

iv Contents
20 Renal Parenchymal Neoplasms 329 31 Disorders of the Adrenal Glands 509

Anobel Y. Odisho, MD, MPH; & Michelle L. McDonald, MD; &
Kirsten L. Greene, MD, MS Christopher J. Kane, MD, FACS
21 Cancer of the Prostate Gland 351 32 Disorders of the Kidneys 521

Matthew R. Cooperberg, MD, MPH; David B. Bayne, MD, MPH;
Samuel L. Washington III, MD; & Jack W. McAninch, MD, FACS, FRCS(E)(Hon); &
Peter R. Carroll, MD, MPH Thomas Chi, MD
22 Genital Tumors 377 33 Diagnosis of Medical Renal Diseases 539

Sima P. Porten, MD, MPH; & Brian K. Lee, MD; & Flavio G. Vincenti, MD
Joseph C. Presti, Jr., MD
34 Acute Kidney Injury and Oliguria 551
23 Urinary Diversion and
Brian K. Lee, MD; & Flavio G. Vincenti, MD
Bladder Substitutions 391
Maxwell V. Meng, MD, MPH; 35 Chronic Kidney Disease and
Susan Barbour, RN, MS, WOCN; & Renal Replacement Therapy 557
Peter R. Carroll, MD, MPH
Brian K. Lee, MD; &
Flavio G. Vincenti, MD
24 Systemic Therapy of Urologic Tumors 407
Vadim S. Koshkin, MD; & Eric J. Small, MD 36 Renal Transplantation 563
John M. Barry, MD
25 Immunotherapy in
Urologic Malignancies 415
37 Disorders of the Ureter and
Arpita Desai, MD; & Eric J. Small, MD Ureteropelvic Junction 571
Barry A. Kogan, MD
26 Radiotherapy of
Urologic Tumors 421
38 Disorders of the Bladder, Prostate,
Yun Rose Li, MD, PhD; and Seminal Vesicles 585
Alexander R. Gottschalk, MD, PhD; &
Mack Roach III, MD Samuel L. Washington III, MD; &
Katsuto Shinohara, MD
27 Neurophysiology and Pharmacology
of the Lower Urinary Tract 453 39 Male Sexual Dysfunction 605
Karl-Erik Andersson, MD, PhD Amanda B. Reed-Maldonado, MD, FACS; &
Tom F. Lue, MD
28 Neurogenic Bladder 473
40 Women’s Sexual Health 631
Anne M. Suskind, MD, MS, FACS
Alan W. Shindel, MD, MAS; &
Tami S. Rowen, MD, MS
29 Urodynamics 485
Anne M. Suskind, MD, MS, FACS 41 Disorders of the Penis and
Male Urethra 645
30 Urinary Incontinence 499
Benjamin N. Breyer, MD, MAS, FACS; &
Tom F. Lue, MD, FACS, ScD (Hon); & Jack W. McAninch, MD, FACS, FRCS(E)(Hon)
Emil A. Tanagho, MD

Contents v
42 Disorders of the Female Urethra 659 46 Genital Gender-Affirming Surgery:

Patient Care, Decision Making, and
Donna Y. Deng, MD, MS
Surgery Options 747
43 Disorders of Sex Development 671 Maurice M. Garcia, MD, MAS
Laurence S. Baskin, MD
47 History and Physical Examination in
Pediatric Urology 769
44 Male Infertility 703
Michael DiSandro, MD
Thomas J. Walsh, MD, MS; &
James F. Smith, MD, MS
48 Introduction to Clinical
Research Design 781
45 The Aging Male 735
June M. Chan, ScD; David Tat, DO; &
James F. Smith, MD, MS; Stacey Kenfield, ScD
Bogdana Schmidt, MD, MPH; &
Thomas J. Walsh, MD, MS Index 793

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Contributors
Karl-Erik Andersson, MD, PhD June M. Chan, ScD
Institute for Regenerative Medicine Program Director, Genitourinary Cancer Epidemiology and
Wake Forest University School of Medicine Population Sciences
Winston Salem, North Carolina Department of Urology
UCSF School of Medicine
Susan Barbour, RN, MS, WOCN San Francisco, California
Palliative Care Services
UCSF School of Medicine Thomas Chi, MD
San Francisco, California Associate Professor and Katzman Endowed Professor in
Clinical Urology
John M. Barry, MD Department of Urology
Professor of Urology and Professor of Surgery UCSF School of Medicine
Division of Abdominal Organ Transplantation San Francisco, California
Organ Health and Science University
Portland, Oregon Matthew R. Cooperberg, MD, MPH
Associate Professor
Laurence S. Baskin, MD Department of Urology
Chief of Pediatric Urology Helen Diller Family Comprehensive Cancer Center
University of California Children’s Medical Center UCSF School of Medicine
UCSF School of Medicine San Francisco, California
Attending Urologist Hillary L. Copp, MD, MS
Children’s Hospital Oakland Associate Professor of Urology and Pediatric Urology
Oakland, California Fellowship Director
Benioff Children’s Hospital
David B. Bayne, MD, MPH UCSF School of Medicine
Endourology Fellow San Francisco, California
UCSF School of Medicine Donna Y. Deng, MD, MS
San Francisco, California Neurourology Lead, Kaiser Permanente Northern
California
Benjamin N. Breyer, MD, MAS, FACS Medical Director, Kaiser NorCal Regional Spina Bifida
Associate Professor and Vice Chair Program
Department of Urology Associate Fellowship Director, Female Pelvic Medicine
UCSF School of Medicine Reconstructive Surgery, Kaiser East Bay/UCSF
San Francisco, California Oakland, California
Peter R. Carroll, MD, MPH Arpita Desai, MD

Professor Clinical Instructor
Ken and Donna Derr-Chevron Endowed Chair in Department of Genitourinary Medical Oncology
Prostate Cancer Helen Diller Family Comprehensive Cancer Center
Department of Urology UCSF School of Medicine
UCSF School of Medicine San Francisco, California
vii

viii Contributors
Michael DiSandro, MD Kirsten L. Greene, MD, MS

Professor of Urology Professor and Chair
Department of Urology Department of Urology
UCSF School of Medicine University of Virginia
San Francisco, California Charlottesville, Virginia
Daniela Franz, MD Hedvig Hricak, MD, PhD

Department of Diagnostic and Interventional Radiology Chair
Klinikum rechts der Isar Department of Radiology
Munich Technical University Memorial Sloan-Kettering Cancer Center
Munich, Germany Professor of Radiology
Cornell University
Thomas W. Gaither, MD, MAS New York, New York
Urology resident
University of California Christopher J. Kane, MD, FACS
Los Angeles, California Dean of Clinical Affairs
UC San Diego School of Medicine
Maurice M. Garcia, MD, MAS CEO, UC San Diego Health Physician Group
Associate Professor of Urology and Anatomy (Adjunct) La Jolla, California
Departments of Urology and Anatomy
UCSF Medical Center Stacey A. Kenfield, ScD
San Francisco, California Associate Professor
Director, Cedars-Sinai Transgender Surgery and Department of Urology
Health Program UCSF School of Medicine
Division of Urology San Francisco, California
Cedars-Sinai Medical Center
Los Angeles, California Barry A. Kogan, MD
Professor, Surgery and Pediatrics
Scott Gerst, MD Falk Chair in Urology
Associate Attending Physician Albany Medical College
Department of Radiology Albany, New York
Memorial Hospital, Memorial Sloane-Kettering
Cancer Center Ryan Kohlbrenner, MD
New York, New York Assistant Professor of Interventional Radiology
Departments of Radiology and Biomedical Imaging
Rolf Gillitzer, MD UCSF School of Medicine
Clinical Director San Francisco, California
Johannes Gutenberg University Medical Center Mainz Badrinath R. Konety, MD, MBA
Mainz, Germany Associate Dean for Innovation
Roy L. Gordon, MD Director of the Institute for Prostate and Urologic Cancers
Professor of Interventional Radiology University of Iowa
Department of Radiology Iowa City, Iowa
San Francisco, California Vadim S. Koshkin, MD
Assistant Clinical Professor
Alexander R. Gottschalk, MD, PhD Genitourinary Medical Oncologist
Professor of Radiation Oncology Departments of Hematology and Oncology
Director of CyberKnife UCSF School of Medicine
Departments of Radiation and Oncology San Francisco, California

Contributors ix
John N. Krieger, MD Anobel Y. Odisho, MD, MPH

Professor of Urology Assistant Professor
University of Washington School of Medicine Department of Urologic Oncology
Seattle, Washington UCSF School of Medicine
Brian K. Lee, MD
Professor of Medicine Sima P. Porten, MD, MPH
The Connie Frank Kidney Transplant Center Assistant professor
UCSF School of Medicine Department of Urology
San Francisco, California UCSF School of Medicine
Yun Rose Li, MD, PhD
Resident Physician Joseph C. Presti, Jr., MD
Departments of Radiation and Oncology Lead for Urologic Oncology
UCSF School of Medicine Kaiser Permanente Northern California
San Francisco, California Oakland, California
Tom F. Lue, MD, FACS, ScD (Hon) Amanda B. Reed-Maldonado, MD, FACS
Professor of Urology Chief, Male Reproductive Urology
Emil Tanagho Endowed Chair in Clinical Urology Department of Urology
Department of Urology Tripler Army Medical Center
UCSF School of Medicine Honolulu, Hawaii
Mack Roach III, MD
Kristin Madden, PharmD Professor of Radiation Oncology and Urology
Pharmacist Department of Urology
Department of Veterans Affairs UCSF School of Medicine
San Antonio, Texas San Francisco Comprehensive Cancer Center
Jack W. McAninch, MD, FACS, FRCS(E)(Hon)
Professor of Urology Tami S. Rowen, MD, MS
UCSF School of Medicine Assistant Professor
San Francisco, California Departments of Obstetrics, Gynecology, and
Reproductive Sciences
Michelle L. McDonald, MD UCSF School of Medicine
Urologist San Francisco, California
San Diego, California
Bogdana Schmidt, MD, MPH
Maxwell V. Meng, MD, MPH Urologic Oncology Fellow
Professor Stanford University Medical Center
Department of Urology Stanford, California
San Francisco, California Alan W. Shindel, MD, MAS
Associate Professor
Hiep T. Nguyen, MD Department of Urology
Associate Professor University of California
Surgery and Urology Davis, California
Harvard Medical School and Children’s Hospital
Boston, Massachusetts

x Contributors
Katsuto Shinohara, MD David Tat, DO

Professor Infectious Disease Specialist
Helen Diller Family Chair in Clinical Urology Moses H. Cone Memorial Hospital
Department of Urology Greensboro, North Carolina
San Francisco, California Joachim W. Thüroff, MD
Professor
Eric J. Small, MD Department of Urology
Professor of Medicine and Urology University Medical Center
Urologic Oncology Program and Program Member, Mannheim, Germany
Comprehensive Cancer Center
UCSF School of Medicine Flavio G. Vincenti, MD
San Francisco, California Professor of Medicine
The Connie Frank Kidney Transplant Center
James F. Smith, MD, MS UCSF School of Medicine
Associate Professor San Francisco, California
Director, Male Reproductive Health
Departments of Urology, Obstetrics, Gynecology, and Thomas J. Walsh, MD, MS
Reproductive Sciences Associate Professor
UCSF School of Medicine Department of Urology
San Francisco, California University of Washington School of Medicine
Seattle, Washington
Professor of Urology Mary K. Wang, MD
Department of Urology Childrens’ Urology
UCSF School of Medicine Austin, Texas
Samuel L. Washington, III, MD
Anne M. Suskind, MD, MS, FACS Urologic Oncology Clinical Fellow
Associate Professor of Urology, Obstetrics, Gynecology, and Department of Urology
Reproductive Sciences UCSF School of Medicine
Director, Neurourology, Female Pelvic Medicine & San Francisco, California
Reconstructive Surgery
UCSF School of Medicine J. Stuart Wolf, Jr., MD, FACS
San Francisco, California Professor, Department of Surgery and Perioperative Care
Dell Medical School
Emil A. Tanagho, MD The University of Texas at Austin
Professor of Urology Austin, Texas

Preface
Smith & Tanagho’s General Urology, nineteenth edition, provides the updated information for the understanding, diagnosis,
and treatment of urological diseases in a concise and well-organized format. The book is up-to-date, to the point, and readable.
Medical students will find this book useful because of its concise, easy-to-follow format, and its breadth of information on
common urological diseases. Residents, as well as practicing physicians in urology, family practice, or general medicine, will find
it an efficient and current reference, particularly because of its emphasis on diagnosis and treatment.
This nineteenth edition has been thoroughly updated with clinical information and current references. The reader will find
that this edition is written in an uncomplicated, straightforward manner that provides relevant clinical information and guide-
lines for diagnosis and management of urologic conditions. Chapters on immunotherapy in urologic malignancies, radiotherapy
of urologic tumors, urinary incontinence, and vascular interventional radiology have all undergone extensive revision. For
this current edition, we have added two chapters on the timely topic of gender dysphoria and introduction to clinical research
design.
Many illustrations and figures have been modernized and improved with added color. The classic fine anatomic drawings
demonstrate well the important clinical findings.
This book has been one of the leading sources of information for students, trainees, and urologists around the world. In addi-
tion to English, this book has been published in many other foreign languages, like Chinese, French, Greek, Italian, Japanese,
Korean, Portuguese, Russian, Spanish, and Turkish.
We greatly appreciate the patience and efforts of our McGraw-Hill staff, the expertise of our contributors, and the support
of our readers.
Jack W. McAninch, MD, FACS, FRCS(E) (Hon)

San Francisco, California, January 2020
xi

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1
1
Anatomy of the
Genitourinary Tract
Emil A. Tanagho, MD; & Tom F. Lue, MD, ScD (Hon), FACS
Urology deals with diseases and disorders of the adrenal ▶▶Blood Supply
gland, the male genitourinary tract, and the female
urinary tract. These systems are illustrated in Figures 1–1 A. Arterial
and 1–2. Each adrenal gland receives three arteries: one from the infe-
rior phrenic artery, one from the aorta, and one from the
ADRENALS renal artery.
B. Venous
▶▶Gross Appearance
A. Anatomy Blood from the right adrenal gland is drained by a very short
vein into the vena cava; the left adrenal vein terminates in the
Each kidney is capped by an adrenal gland, and both left renal vein.
organs are enclosed within Gerota’s (perirenal) fascia.
Each adrenal gland weighs 4–5 g. The right adrenal is tri- ▶▶Lymphatics
angular in shape; the left is more rounded and crescentic.
The average dimensions are 3 cm width, 5 cm length, and The lymphatic vessels accompany the suprarenal vein and
1 cm thickness. Each gland is composed of a cortex, chiefly drain into the lumbar lymph nodes.
influenced by the pituitary gland, and a medulla derived
from chromaffin tissue (Avisse et al, 2000; O’Donoghue KIDNEYS
et al, 2010).
B. Relations A. Anatomy
Figure 1–2 shows the relationships between the adrenals and The kidneys lie along the borders of the psoas muscles and
other organs. The right adrenal lies between the liver and are therefore obliquely placed. The position of the liver
the vena cava. The left adrenal lies close to the aorta and is causes the right kidney to be lower than the left (Figures 1–2
covered on its lower surface by the pancreas. The spleen lies and 1–3). The adult kidney weighs between 125 and 170 g in
superior and lateral to it. men and 115 and 155 g in women. It is about 10–12 cm long,
5–7 cm wide, and 3–5 cm thick.
The kidneys are supported by the perirenal fat (which is
▶▶Histology enclosed in the perirenal fascia), the renal vascular pedicle,
The adrenal cortex, which makes up 85% of the mass, is com- abdominal muscle tone, and the general bulk of the abdomi-
posed of three distinct layers: the outer zona glomerulosa, the nal viscera (Rusinek et al, 2004). Variations in these factors
middle zona fasciculata, and the inner zona reticularis. The permit variations in the degree of renal mobility. The aver-
medulla lies centrally and is made up of polyhedral cells with age descent on inspiration or on assuming the upright posi-
hormone-containing granular cytoplasm. These chromaf- tion is 4–5 cm. Lack of mobility suggests abnormal fixation
fin cells are accompanied by a small number of sympathetic (eg, perinephritis), but extreme mobility is not necessarily
ganglion cells. pathologic.
McAninch_CH01_p001-p016.indd 1 07/02/20 9:58 AM

2 SMITH & TANAGHO’S GENERAL UROLOGY
▲▲Figure 1–1. Anatomy of the male genitourinary tract. The upper tract and midtract have urologic function only.
The lower tract has both genital and urinary functions.

ANATOMY OF THE GENITOURINARY TRACT CHAPTER 1 3
▲▲Figure 1–2. Relations between the kidneys, ureters, and bladder (anterior aspect).
On longitudinal section (Figure 1–4), the kidney is seen gastrointestinal symptoms that accompany kidney diseases
to be made up of an outer cortex, a central medulla, and the (Glassberg, 2002).
internal calices and pelvis. The cortex is homogeneous in
appearance. Portions of it project toward the pelvis between ▶▶Histology
the papillae and fornices and are called the columns of A. Nephron
Bertin. The medulla consists of numerous pyramids formed
by the converging collecting renal tubules, which drain into The functioning unit of the kidney is the nephron, which is
the minor calices at the tip of the papillae. composed of a tubule that has both secretory and excretory
functions (Figure 1–4). The secretory portion is contained
largely within the cortex and consists of a renal corpuscle and
B. Relations
the secretory part of the renal tubule. The excretory portion of
Figures 1–2 and 1–3 show the relationships between the this duct lies in the medulla. The renal corpuscle is composed
kidneys and adjacent organs and structures. Their intimacy of the vascular glomerulus, which projects into Bowman’s cap-
with intraperitoneal organs and the autonomic innervation sule, which, in turn, is continuous with the epithelium of the
that they share with these organs explain, in part, some of the proximal convoluted tubule. The secretory portion of the renal

▲▲Figure 1–3. Relations between the kidneys (posterior aspect). The dashed lines represent the outline of the kidneys,
where they are obscured by overlying structures.
tubule is made up of the proximal convoluted tubule, the loop of the posterior surface. The anterior branch supplies both
of Henle, and the distal convoluted tubule. upper and lower poles as well as the entire anterior surface.
The excretory portion of the nephron is the collecting The renal arteries are all end arteries.
tubule, which is continuous with the distal end of the ascend- The renal artery branches further divide into interlobar
ing limb of the convoluted tubule. It empties its contents arteries, which travel in the columns of Bertin (between the
through the tip (papilla) of a pyramid into a minor calyx. pyramids) and then arch along the base of the pyramids
(arcuate arteries). These arteries then divide as interlobular
B. Supporting Tissue arteries. From these vessels, smaller (afferent) branches pass
to the glomeruli. From the glomerular tuft, efferent arterioles
The renal stroma is composed of loose connective tissue and
pass to the tubules in the stroma.
contains blood vessels, capillaries, nerves, and lymphatics.
B. Venous
▶▶Blood Supply (Figures 1–2, 1–4, and 1–5)
The renal veins are paired with the arteries, but any of them
A. Arterial
will drain the entire kidney if the others are tied off.
Usually there is one renal artery, a branch of the aorta that Although the renal artery and vein are usually the sole blood
enters the hilum of the kidney between the pelvis, which vessels of the kidney, accessory renal vessels are common and
normally lies posteriorly, and the renal vein. It may branch may be of clinical importance if they are so placed so as to com-
before it reaches the kidney, and two or more separate arter- press the ureter, in which case hydronephrosis may result.
ies may be noted (Budhiraja et al, 2010). In duplication of the
pelvis and ureter, it is common for each renal segment to have
its own arterial supply. ▶▶Nerve Supply
The renal artery divides into anterior and posterior The renal nerves derived from the renal plexus accompany
branches. The posterior branch supplies the midsegment the renal vessels throughout the renal parenchyma.

▲▲Figure 1–4. Anatomy and histology of the kidney and ureter. Upper left: Diagram of the nephron and its blood supply.
(Courtesy of Merck, Sharp, Dohme: Seminar. 1947; 9[3].) Upper right: Cast of the pelvic caliceal system and the arterial
supply of the kidney. Middle: Renal calices, pelvis, and ureter (posterior aspect). Lower left: Histology of the ureter. The
smooth-muscle bundles are arranged in both spirally and longitudinally. Lower right: Longitudinal section of kidney
showing calices, pelvis, ureter, and renal blood supply (posterior aspect).

▲▲Figure 1–5. (A) The posterior branch of the renal artery and its distribution to the central segment of the posterior
surface of the kidney. (B) Branches of the anterior division of the renal artery supplying the entire anterior surface of the
kidney as well as the upper and lower poles at both surfaces. The segmental branches lead to interlobar, arcuate, and
interlobular arteries. (C) The lateral convex margin of the kidney. Brödel’s line, which is 1 cm from the convex margin, is
the bloodless plane demarcated by the distribution of the posterior branch of the renal artery.

▶▶Lymphatics
The lymphatics of the kidney drain into the lumbar lymph
nodes.
CALICES, RENAL PELVIS, AND URETER
A. Anatomy
1. Calices—The tips of the minor calices (8–12 in number)

are indented by the projecting pyramids (Figure 1–4). These
calices unite to form two or three major calices that join to
form the renal pelvis (Sozen et al, 2008).
2. Renal pelvis—The pelvis may be entirely intrarenal or
partly intrarenal and partly extrarenal. Inferomedially, it
tapers to join the ureter.
▲▲Figure 1–6. Anatomy and relations between the
3. Ureter—The adult ureter is about 30 cm long, varying ureters, bladder, prostate, seminal vesicles, and vasa
in direct relation to the height of the individual. It follows a deferentia (anterior view).
rather smooth S curve. Areas that stones are often impacted
are (a) at the ureteropelvic junction, (b) where the ureter
crosses over the iliac vessels, and (c) where it courses through helical and longitudinal smooth-muscle fibers. They are not
the bladder wall. arranged in discrete layers. The outermost adventitial coat is
composed of fibrous connective tissue.
B. Relations
▶▶Blood Supply
1. Calices—The calices are intrarenal and are intimately A. Arterial
related to the renal parenchyma.
The renal calices, pelvis, and upper ureters derive their blood
2. Renal pelvis—If the pelvis is partly extrarenal, it lies along supply from the renal arteries; the midureter is fed by the
the lateral border of the psoas muscle and on the quadratus internal spermatic (or ovarian) arteries. The lowermost por-
lumborum muscle; the renal vascular pedicle is just anterior tion of the ureter is served by branches from the common
to it. The left renal pelvis lies at the level of the first or second iliac, internal iliac (hypogastric), and vesical arteries.
lumbar vertebra; the right pelvis is a little lower.
3. Ureter—On their course downward, the ureters lie on the B. Venous
psoas muscles, pass medially to the sacroiliac joints, and then The veins of the renal calices, pelvis, and ureters are paired
swing laterally near the ischial spines before passing medi- with the arteries.
ally to enter the base of the bladder (Figure 1–2). In females,
the uterine arteries are closely related to the juxtavesical por-
tion of the ureters. The ureters are covered by the posterior
▶▶Lymphatics
peritoneum; their lowermost portions are closely attached to The lymphatics of the upper portions of the ureters as well
it, while the juxtavesical portions are embedded in vascular as those from the pelvis and calices enter the lumbar lymph
retroperitoneal fat (Koff, 2008). nodes. The lymphatics of the midureter pass to the internal
The vasa deferentia, as they leave the internal inguinal iliac (hypogastric) and common iliac lymph nodes; the lower
rings, sweep over the lateral pelvic walls anterior to the ureteral lymphatics empty into the vesical and hypogastric
ureters (Figure 1–6). They lie medial to the latter before join- lymph nodes.
ing the seminal vesicle and penetrating the base of the pros-
tate to become the ejaculatory ducts. BLADDER
▶▶Histology (Figure 1–4) ▶▶Gross Appearance

The walls of the calices, pelvis, and ureters are composed of The bladder is a hollow muscular organ that serves as a res-
transitional cell epithelium under which lies loose connec- ervoir for urine. In women, its posterior wall and dome are
tive tissue (lamina propria). External to these are a mixture of invaginated by the uterus. The adult bladder normally has

a capacity of 400–500 mL. The wall of the bladder is about ▶▶Blood Supply
3–5 mm in thickness; it is thinner when it is distended.
A. Arterial
A. Anatomy The bladder is supplied by the superior, middle, and inferior
When empty, the adult bladder lies behind the pubic sym- vesical arteries, which arise from the anterior trunk of the
physis and is largely a pelvic organ. In infants and children, internal iliac (hypogastric) artery, and by smaller branches
it is situated higher (Berrocal et al, 2002). When it is full, it from the obturator and inferior gluteal arteries. In females,
rises well above the symphysis and can readily be palpated the uterine and vaginal arteries also send branches to the
or percussed. When overdistended, as in acute or chronic bladder.
urinary retention, it may cause the lower abdomen to bulge
visibly. B. Venous
Extending from the dome of the bladder to the umbilicus Surrounding the bladder is a rich plexus of veins that ulti-
is a fibrous cord, the median umbilical ligament, which rep- mately empties into the internal iliac (hypogastric) veins.
resents the obliterated urachus. The ureters enter the bladder
posteroinferiorly in an oblique manner and at these points ▶▶Nerve Supply
are about 5 cm apart (Figure 1–6). The orifices, situated at
the extremities of the crescent-shaped interureteric ridge that The bladder receives innervation from sympathetic and
forms the proximal border of the trigone, are about 2.5 cm parasympathetic nervous systems. The sensory afferent of
apart. The trigone occupies the area between the ridge and the bladder originates from both subepithelial nerve endings
the bladder neck. and nerve fibers between detrusor muscle bundles (Andersson,
The internal sphincter, or bladder neck, is not a true cir- 2010; Birder et al, 2010; McCloskey, 2010).
cular sphincter but a thickening formed by interlaced and
converging muscle fibers of the detrusor as they pass distally ▶▶Lymphatics
to become the smooth muscle component of the urethra. The lymphatics of the bladder drain into the vesical, external
iliac, internal iliac (hypogastric), and common iliac lymph
B. Relations nodes.
In males, the bladder is related posteriorly to the seminal
vesicles, vasa deferentia, ureters, and rectum (Figures 1–7 PROSTATE GLAND
and 1–8). In females, the uterus and vagina are interposed
between the bladder and rectum (Figure 1–9). The dome ▶▶Gross Appearance
and posterior surfaces are covered by peritoneum; hence, in
A. Anatomy
this area, the bladder is closely related to the small intestine
and sigmoid colon. In both males and females, the bladder is The prostate is a fibromuscular and glandular organ lying just
related to the posterior surface of the pubic symphysis, and, inferior to the bladder (Figures 1–6 and 1–7). The normal
when distended, it is in contact with the lower abdominal prostate weighs about 20 g and contains the posterior urethra,
wall. which is about 2.5 cm in length. It is supported anteriorly by
the puboprostatic ligaments and inferiorly by the urogenital
diaphragm (Figure 1–6). The prostate is perforated posteri-
▶▶Histology (Figure 1–10) orly by the ejaculatory ducts, which pass obliquely to empty
The mucosa of the bladder is composed of transitional epi- through the verumontanum on the floor of the prostatic ure-
thelium. Beneath it is a well-developed submucosal layer thra just proximal to the striated external urinary sphincter
formed largely of connective and elastic tissues. The mucosa (Figure 1–11).
may be considered as a single functional unit that consists The prostate can be subdivided into two ways: by lobe
of the epithelial layer, basement membrane, and lamina pro- or by zone. The lobe classification is often used in cystoure-
pria. Physical or chemical stress on the bladder elicits releases throscopic examinations and consists of five lobes: anterior,
of multiple factors that modulate afferent and efferent nerve posterior, median, right lateral, and left lateral. The zone clas-
activities (Fry and Vahabi, 2016). External to the submu- sification is often used in pathology. McNeal (1981) divides
cosa is the detrusor muscle that is made up of a mixture of the prostate into four zones: peripheral zone, central zone
smooth-muscle fibers arranged at random in a longitudi- (surrounds the ejaculatory ducts), transitional zone (sur-
nal, circular, and spiral manner without any layer formation rounds the urethra), and anterior fibromuscular zone (Myers
or specific orientation except for proximity to the internal et al, 2010) (Figure 1–12). The segment of urethra that tra-
meatus, where the detrusor muscle assumes three definite verses the prostate gland is the prostatic urethra. It is lined
layers: inner longitudinal, middle circular, and outer longitu- by an inner longitudinal layer of muscle (continuous with
dinal (John et al, 2001). a similar layer of the vesical wall). Incorporated within the

▲▲Figure 1–7. (A) Anatomic relationship between the bladder, prostate, prostatomembranous urethra, and root of
the penis. (B) Histology of the testis. Seminiferous tubules lined by supporting basement membrane for the Sertoli and
spermatogenic cells. The latter are in various stages of development. (C) Cross sections of the testis and epididymis.
(Images [A] and [C] reproduced with permission from Walsh PC, Campbell MF: Campbell’s Urology, 6th ed. Philadelphia, PA:
Saunders; 1992.)

▲▲Figure 1–8. Top: Relations between the bladder, prostate, seminal vesicles, penis, urethra, and scrotal contents.
Lower left: Transverse section through the penis. The paired upper structures are the corpora cavernosa. The single
lower body surrounding the urethra is the corpus spongiosum. Lower right: Fascial planes of the lower genitourinary
tract. (After Wesson.)
prostate gland is an abundant amount of smooth muscula- separated from the rectum by the two layers of Denonvilliers’
ture derived primarily from the external longitudinal bladder fascia, serosal rudiments of the pouch of Douglas, which
musculature. This musculature represents the involuntary once extended to the urogenital diaphragm (Raychaudhuri
smooth muscle sphincter of the posterior urethra in males. and Cahill, 2008) (Figure 1–8).
B. Relations ▶▶Histology (Figure 1–10)

The prostate gland lies behind the pubic symphysis. Located The prostate consists of a thin fibrous capsule under
closely to the posterosuperior surface are the vasa deferentia which lie circularly oriented smooth-muscle fibers and
and seminal vesicles (Figure 1–7). Posteriorly, the prostate is collagenous tissue that surrounds the urethra (involuntary

▲▲Figure 1–9. Anatomy and relations of the bladder, urethra, uterus and ovary, vagina, and rectum.
▲▲Figure 1–10. Left: Histology of the prostate. Epithelial glands embedded in a mixture of connective and elastic tissue
and smooth muscle. Right: Histology of the bladder. The mucosa is transitional cell in type and lies on a well-developed
submucosal layer of connective tissue. The detrusor muscle is composed of interlacing longitudinal, circular, and spiral
smooth-muscle bundles.

▲▲Figure 1–11. Section of the prostate gland shows the

prostatic urethra, verumontanum, and crista urethralis,
in addition to the opening of the prostatic utricle and
the two ejaculatory ducts in the midline. Note that the
prostate is surrounded by the prostatic capsule, which
is covered by another prostatic sheath derived from the
endopelvic fascia. The prostate is resting on the genitouri-
nary diaphragm. (Reproduced with permission from Walsh
PC, Campbell MF: Campbell’s Urology, 6th ed. Philadelphia,
PA: Saunders; 1992.) ▲▲Figure 1–12. Anatomy of the prostate gland. Prostatic
adenoma develops from the periurethral glands at the
site of the median or lateral lobes. The posterior lobe,
however, is prone to cancerous degeneration. (Adapted
sphincter). Deep in this layer lies the prostatic stroma,
with permission from McNeal JE: The zonal anatomy of the
composed of connective tissues and smooth-muscle fibers
prostate. Prostate 1981;2(1):35–49.)
in which are embedded the epithelial glands. These glands
drain into the major excretory ducts (about 25 in number),
which open chiefly on the floor of the urethra between
the verumontanum and the vesical neck. Just beneath ▶▶Lymphatics
the transitional epithelium of the prostatic urethra lie the The lymphatics from the prostate drain into the internal iliac
periurethral glands. (hypogastric), sacral, vesical, and external iliac lymph nodes
(Saokar et al, 2010).
▶▶Blood Supply
A. Arterial SEMINAL VESICLES
The arterial supply to the prostate is derived from the inferior
vesical, internal pudendal, and middle rectal (hemorrhoidal) ▶▶Gross Appearance
arteries. The seminal vesicles lie just cephalic to the prostate under
the base of the bladder (Figures 1–6 and 1–7). They are about
B. Venous 6 cm long and quite soft. Each vesicle joins its corresponding
vas deferens to form the ejaculatory duct (Kim et al, 2009).
The veins from the prostate drain into the periprostatic The ureters lie medial to each, and the rectum is contiguous
plexus, which has connections with the deep dorsal vein of with their posterior surfaces.
the penis and the internal iliac (hypogastric) veins.
▶▶Histology
▶▶Nerve Supply The mucous membrane is pseudostratified. The submu-
The prostate gland receives a rich innervation from the sym- cosa consists of dense connective tissue covered by a thin
pathetic and parasympathetic nerves of the inferior hypogas- layer of muscle that, in turn, is encapsulated by connective
tric plexus. tissue.

▶▶Blood Supply EPIDIDYMIS

The blood supply of the seminal vesicles is similar to that of
the prostate gland. ▶▶Gross Appearance
A. Anatomy
▶▶Nerve Supply The upper portion of the epididymis (globus major) is con-
The nerve supply is mainly from the sympathetic nerve nected to the testis by numerous efferent ducts from the testis
plexus. (Figure 1–7). The epididymis consists of a markedly coiled
duct that, at its lower pole (globus minor), is continuous with
▶▶Lymphatics the vas deferens. An appendix of the epididymis is often seen
on its upper pole; this is a cystic body that in some cases is
The lymphatics of the seminal vesicles are those that serve
pedunculated, but in others, it is sessile.
the prostate.
B. Relations
SPERMATIC CORD
The epididymis lies posterolateral to the testis and is nearest
▶▶Gross Appearance to the testis at its upper pole. Its lower pole is connected to
the testis by fibrous tissue. The vas lies posteromedial to the
The two spermatic cords extend from the internal ingui- epididymis.
nal rings through the inguinal canals to the testicles
(Figure 1–7). Each cord contains the vas deferens, the
internal and external spermatic arteries, the artery of
▶▶Histology
the vas, the venous pampiniform plexus (which forms The epididymis is covered by serosa. The ductus epididy-
the spermatic vein superiorly), lymph vessels, and nerves midis is lined by pseudostratified columnar epithelium
(Jen et al, 1999). The entire cord contents are enclosed in throughout its length.
investing layers of thin fascia. A few fibers of the cremaster
muscle insert on the cords in the inguinal canal (Bhosale ▶▶Blood Supply
et al, 2008; Kim et al, 2009). A. Arterial
▶▶Histology The arterial supply to the epididymis comes from the internal
spermatic artery and the artery of the vas (deferential artery).
The fascia covering the cord is formed of loose connective
tissue that supports arteries, veins, nerve, and lymphatics. B. Venous
The vas deferens is a small, thick-walled tube consisting of an
internal mucosa and submucosa surrounded by three well- The venous blood drains into the pampiniform plexus, which
defined layers of smooth muscle encased in a covering of becomes the spermatic vein.
fibrous tissue. Above the testes, this tube is straight. Its proxi-
mal 4 cm tends to be convoluted. ▶▶Lymphatics
The lymphatics drain into the external iliac and internal iliac
▶▶Blood Supply (hypogastric) lymph nodes.
A. Arterial
TESTIS
The external spermatic artery, a branch of the inferior epigas-
tric, supplies the fascial coverings of the cord. The internal ▶▶Gross Appearance
spermatic artery passes through the cord on its way to the
testis. The deferential artery is close to the vas. A. Anatomy
The average testicle measures about 4 × 3 × 2.5 cm
B. Venous (Figure 1–7). The volume can be measured by an orchidom-
The veins from the testis and the coverings of the spermatic eter or by a formula with ultrasonic measurement (length ×
cord form the pampiniform plexus, which, at the internal width × height × 0.71). The average volume is 18 mL (rang-
inguinal ring, unites to form the spermatic vein. ing from 12 to 30 mL). The testicle has a dense fascial cov-
ering called the tunica albuginea testis, which, posteriorly,
is invaginated somewhat into the body of the testis to form
▶▶Lymphatics the mediastinum testis. This fibrous mediastinum sends
The lymphatics from the spermatic cord empty into the fibrous septa into the testis, thus separating it into about
external iliac lymph nodes. 250 lobules.

The testis is covered anteriorly and laterally by the visceral SCROTUM

layer of the serous tunica vaginalis, which is continuous with
the parietal layer that separates the testis from the scrotal ▶▶Gross Appearance
wall (Bidarkar and Hutson, 2005). A small amount of fluid
normally exists within the tunica vaginalis sac. At the upper Beneath the corrugated skin of the scrotum lies the dartos
pole of the testis is the appendix testis, a small pedunculated muscle. Deep to this are the three fascial layers derived from
or sessile body similar in appearance to the appendix of the the abdominal wall at the time of testicular descent. Beneath
epididymis. these is the parietal layer of the tunica vaginalis (Kim et al,
2007).
B. Relations The scrotum is divided into two sacs by a septum of con-
nective tissue. The scrotum not only supports the testes but
The testis is closely attached posterolaterally to the epididy- also, by relaxation or contraction of its muscular layer, helps
mis, particularly at its upper and lower poles (Klonisch et al, to regulate their temperature.
2004).
▶▶Histology
▶▶Histology (Figure 1–7) The dartos muscle, under the skin of the scrotum, is nonstri-
Each lobule contains one to four markedly convoluted ated. The deeper layer is made up of connective tissue.
seminiferous tubules, each of which is about 60 cm long.
These ducts converge at the mediastinum testis, where ▶▶Blood Supply
they connect with the efferent ducts that drain into the A. Arterial
epididymis.
The seminiferous tubule has a basement membrane con- The arteries to the scrotum arise from the femoral, internal
taining connective and elastic tissue. This supports the semi- pudendal, and inferior epigastric arteries.
niferous cells that are of two types: (1) Sertoli (supporting)
cells and (2) spermatogenic cells. The stroma between the B. Venous
seminiferous tubules contains connective tissue in which the The veins are paired with the arteries.
interstitial Leydig cells are located.
▶▶Lymphatics
▶▶Blood Supply The lymphatics drain into the superficial inguinal and subin-
The blood supply to the testes is closely associated with that guinal lymph nodes.
to the kidneys because of the common embryologic origin of
the two organs. PENIS AND MALE URETHRA
A. Arterial
The arteries to the testes (internal spermatics) arise from the The penis is composed of two corpora cavernosa and the
aorta just below the renal arteries and course through the corpus spongiosum, which contains the urethra. The corpus
spermatic cords to the testes, where they anastomose with spongiosum enlarges distally and forms the glans penis. Each
the arteries of the vasa deferentia that branch off from the corpus is enclosed in a fascial sheath (tunica albuginea), and
internal iliac (hypogastric) artery. all three corpora are surrounded by a thick fibrous envelope
known as Buck’s fascia. A covering of skin, devoid of fat, is
B. Venous loosely wrapped these bodies. The prepuce forms a hood
The blood from the testis returns in the pampiniform plexus over the glans.
of the spermatic cord. At the internal inguinal ring, the Beneath the skin of the penis (and scrotum) and extend-
pampiniform plexus forms the spermatic vein. ing from the base of the glans to the urogenital diaphragm is
The right spermatic vein enters the vena cava just below Colles’ fascia, which is continuous with Scarpa’s fascia of the
the right renal vein; the left spermatic vein empties into the lower abdominal wall (Figure 1–8).
left renal vein. The proximal ends of the corpora cavernosa are attached
to the pelvic bones just anterior to the ischial tuberosities.
The ischiocavernosus muscles insert into the lateral surface
▶▶Lymphatics of the tunica albuginea at the proximal corpora cavernosa.
The lymphatic vessels from the testes pass to the lumbar Occupying a depression of their ventral surface in the midline
lymph nodes, which, in turn, are connected to the medias- is the corpus spongiosum, which is connected proximally to
tinal nodes. the undersurface of the urogenital diaphragm, below which

lies the urethral bulb. This portion of the corpus spongiosum FEMALE URETHRA
is surrounded by the bulbospongiosus muscle.
The suspensory ligament of the penis arises from the linea ▶▶Gross Appearance
alba and pubic symphysis and inserts into the fascial covering
of the corpora cavernosa. The adult female urethra is about 4 cm long and 8 mm in
diameter. It is slightly curved and lies beneath the pubic sym-
physis just anterior to the vagina.
▶▶Histology
A. Corpora and Glans Penis ▶▶Histology
The corpora cavernosa, the corpus spongiosum, and the The epithelial lining of the female urethra is squamous in
glans penis are composed of smooth muscles, intracaverno- its distal portion and pseudostratified or transitional in the
sal struts (corpus cavernosum only), and endothelium-lined remainder. The submucosa is made up of connective and
sinusoids. The sympathetic and parasympathetic (as well as elastic tissues and spongy venous spaces. Embedded in it are
the nonadrenergic, noncholinergic [NANC]) nerve termi- many periurethral glands, which are most numerous distally;
nals are often seen around the vessels and near the smooth the largest of these are the periurethral glands of Skene that
muscles. open on the floor of the urethra just inside the meatus.
External to the submucosa is a longitudinal layer of
B. Urethra smooth muscle continuous with the inner longitudinal
layer of the bladder wall. Surrounding this is a heavy layer
The urethral mucosa that traverses the glans penis is formed
of circular smooth-muscle fibers extending from the exter-
of squamous epithelium. Proximal to this, the mucosa is tran-
nal vesical muscular layer. This constitutes the involuntary
sitional in type. Underneath the mucosa is the submucosa that
internal urethral sphincter. Distal to this is the external stri-
contains connective and elastic tissue and smooth muscle. In
ated (voluntary) sphincter surrounding the middle third of
the submucosa are the numerous glands of Littre, whose ducts
the urethra composed of smooth and striated muscles within
connect with the urethral lumen. The urethra is surrounded by
the midurethra (Ashton-Miller and Delancey, 2009; Morgan
the vascular corpus spongiosum and the glans penis.
et al 2009; Thor and de Groat, 2010).
▶▶Blood Supply ▶▶Blood Supply

A. Arterial The arterial supply to the female urethra is derived from the
The penis and urethra are supplied by the internal pudendal inferior vesical, vaginal, and internal pudendal arteries. Blood
arteries. Each artery divides into a cavernous artery of the from the urethra drains into the internal pudendal veins.
penis (which supplies the corpora cavernosa), a dorsal artery
of the penis, and the bulbourethral artery. These branches ▶▶Lymphatics
supply the corpus spongiosum, the glans penis, and the ure- Lymphatic drainage from the external portion of the urethra
thra. Accessory pudendal arteries originate from inferior is to the inguinal and subinguinal lymph nodes. Drainage
vesical, obturator, or other arteries may also supply the penis from the deep urethra is into the internal iliac (hypogastric)
(Henry et al, 2017). lymph nodes.
B. Venous BIBLIOGRAPHY
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17
2
Embryology of the
Genitourinary System
Emil A. Tanagho, MD; Hiep T. Nguyen, MD;

& Michael DiSandro, MD
At birth, the genital and urinary systems are related only in nearby primary nephric duct as it grows caudally to join
the sense that they share certain common passages. Embryo- the cloaca (Figure 2–1). This primary nephric duct is now
logically, however, they are intimately related. Because of the called the mesonephric duct. After establishing their con-
complex interrelationships of the embryonic phases of the nection with the nephric duct, the primordial tubules elon-
two systems, they are discussed here as five subdivisions: gate and become S-shaped. As the tubules elongate, a series
the nephric system, the vesicourethral unit, the gonads, the of secondary branches increase their surface exposure,
genital duct system, and the external genitalia. thereby enhancing their capacity for interchanging material
with the blood in adjacent capillaries. Leaving the glomeru-
NEPHRIC SYSTEM lus, the blood is carried by one or more efferent vessels that
soon break up into a rich capillary plexus closely related to
The nephric system develops progressively as three distinct
the mesonephric tubules. The mesonephros, which forms
entities: pronephros, mesonephros, and metanephros.
early in the 4th week, reaches its maximum size by the end
of the second month.
▶▶Pronephros
The pronephros is the earliest nephric stage in humans, and ▶▶Metanephros
it corresponds to the mature structure of the most primitive
The metanephros, the final phase of development of the
vertebrate. It extends from the 4th to the 14th somites and
nephric system, originate from both the intermediate meso-
consists of 6–10 pairs of tubules. These open into a pair of
derm and the mesonephric duct. Development begins in the
primary ducts that are formed at the same level, extend cau-
5–6-mm embryo with a budlike outgrowth from the meso-
dally, and eventually reach and open into the cloaca. The pro-
nephric duct as it bends to join the cloaca. This ureteral bud
nephros is a vestigial structure that disappears completely by
grows cephalad and collects mesoderm from the nephro-
the 4th week of embryonic life (Figure 2–1).
genic cord of the intermediate mesoderm around its tip. This
mesoderm with the metanephric cap moves, with the grow-
▶▶Mesonephros ing ureteral bud, more and more cephalad from its point of
The mature excretory organ of the larger fish and amphib- origin. During this cephalic migration, the metanephric cap
ians corresponds to the embryonic mesonephros. It is becomes progressively larger, and rapid internal differentia-
the principal excretory organ during early embryonic life tion takes place. Meanwhile, the cephalic end of the ureteral
(4–8 weeks). It, too, gradually degenerates, although parts bud expands within the growing mass of metanephrogenic
of its duct system become associated with the male repro- tissue to form the renal pelvis (Figure 2–1). Numerous out-
ductive organs. The mesonephric tubules develop from the growths from the renal pelvic dilatation push radially into
intermediate mesoderm caudal to the pronephros shortly this growing mass and form hollow ducts that branch and
before pronephric degeneration. The mesonephric tubules rebranch as they push toward the periphery. These form the
differ from those of the pronephros in that they develop a primary collecting ducts of the kidney. Mesodermal cells
cuplike outgrowth into which a knot of capillaries is pushed. become arranged in small vesicular masses that lie close to
This is called Bowman’s capsule, and the tuft of capillaries the blind end of the collecting ducts. Each of these vesicular
is called a glomerulus. In their growth, the mesonephric masses will form a uriniferous tubule draining into the duct
tubules extend toward and establish a connection with the nearest to its point of origin.

▲▲Figure 2–1. Schematic representation of the development of the nephric system. Only a few of the tubules of the
pronephros are seen early in the 4th week, while the mesonephric tissue differentiates into mesonephric tubules that
progressively join the mesonephric duct. During this time, the first sign of the ureteral bud from the mesonephric duct is
seen. At 6 weeks, the pronephros has completely degenerated and the mesonephric tubules start to do so. The ureteral
bud grows dorsocranially and has met the metanephrogenic cap. At the 8th week, there is cranial migration of the differ-
entiating metanephros. The cranial end of the ureteric bud expands and starts to show multiple successive outgrowths.
(Data from several sources.)
As the kidney grows, increasing numbers of tubules are the ends of the anterior pronephric tubules; (4) this pronephric
formed in its peripheral zone. These vesicular masses develop duct serves subsequently as the mesonephric duct and as such
a central cavity and become S-shaped. One end of the S gives rise to the ureter; (5) the nephric duct reaches the cloaca
coalesces with the terminal portion of the collecting tubules, by independent caudal growth; and (6) the embryonic ureter
resulting in a continuous canal. The proximal portion of the is an outgrowth of the nephric duct, yet the kidney tubules dif-
S develops into the distal and proximal convoluted tubules ferentiate from adjacent metanephric blastema.
and into Henle’s loop; the distal end becomes the glomeru-
lus and Bowman’s capsule. At this stage, the undifferentiated
mesoderm and the immature glomeruli are readily visible on ▶▶Molecular Mechanisms of Renal and
microscopic examination (Figure 2–2). The glomeruli are
Ureteral Development
fully developed by the 36th week or when the fetus weighs The kidney and the collecting system originate from the
2500 g (Osathanondh and Potter, 1964a, b). The metaneph- interaction between the mesonephric duct (Wolffian duct)
ros arises opposite the 28th somite (fourth lumbar segment). and the metanephric mesenchyme (MM). The uretic bud
At term, it has ascended to the level of the first lumbar or (UB) forms as an epithelial outpouching from the meso-
even the twelfth thoracic vertebra. This ascent of the kidney nephric duct and invades the surrounding MM. Reciprocal
is due not only to actual cephalic migration but also to differ- induction between the UB and MM results in branching
ential growth in the caudal part of the body. During the early and elongation of the UB from the collecting system and in
period of ascent (7th–9th weeks), the kidney slides above the condensation and epithelial differentiation of MM around
arterial bifurcation and rotates 90°. Its convex border is now the branched tips of the UB. Branching of the UB occurs
directed laterally, not dorsally. Ascent proceeds more slowly approximately 15 times during human renal development,
until the kidney reaches its final position. generating approximately 300,000 and 1 million nephrons
Certain features of these three phases of development must per kidney (Nyengaard and Bendtsen, 1992).
be emphasized: (1) the three successive units of the system This process of reciprocal induction is dependent on the
develop from the intermediate mesoderm; (2) the tubules at all expression of specific factors. Glial cell-derived neurotrophic
levels appear as independent primordia and only secondarily factor (GDNF) is the primary inducer of ureteric budding
unite with the duct system; (3) the nephric duct is laid down (Costantini and Shakya, 2006). GDNF interacts with sev-
as the duct of the pronephros and develops from the union of eral different proteins from the MM (eg, Wt1, Pax2, Eyal,

EMBRYOLOGY OF THE GENITOURINARY SYSTEM CHAPTER 2 19
▲▲Figure 2–2. Progressive stages in the differentiation of the nephrons and their linkage with the branching collecting
tubules. A small lump of metanephric tissue is associated with each terminal collecting tubule. These are then arranged
in vesicular masses that later differentiate into a uriniferous tubule draining into the duct near which it arises. At one
end, Bowman’s capsule and the glomerulus differentiate; the other end establishes communication with the nearby
collecting tubules.
Six1, Sall1) and from the UB itself (Pax2, Lim1, Ret) result- termination and tubule maintenance (hepatocyte growth fac-
ing in outgrowth of the UB (reviewed by Shah et al, 2004). tor, transforming growth factor-α, epidermal growth factor
Proper activation of the Ret/GDNF signaling pathway in receptor) (reviewed by Shah et al, 2004). BMP7, SHH, and
the tip of UB epithelium appears to be essential in the pro- Wnt11 produced from the branching ureteric bud induce the
gression of branching morphogenesis (reviewed by Michos, MM to differentiate. These factors induce the activation of
2009). B-catenin and Gata3 are important regulators of Ret Pax2, α-8-integrin, and Wnt4 in the renal mesenchymal cells,
expression, and correct activity of Ret is regulated by posi- resulting in condensation of the MM and the formation of
tive (Wnt11 from MM) and negative (Sprouty1 from the UB) pretubular aggregate and primitive renal vesicle (reviewed by
feedback signaling. Additional specific factors are required for Burrow, 2000). With the continued induction from the UB
(1) early branching (eg, Wnt4 and Wnt11, fgf 7–10); (2) late and the autocrine activity of Wnt4, the pretubular aggregates
branching and maturation (bmp2, activin); and (3) branching differentiate into comma-shaped bodies. Platelet-derived

growth factor α-β and vascular endothelial growth factor ureter and is most commonly associated with an accessory
expression are required for initiating the migration of endo- ureteral bud in a duplicated system, but it also can be seen in
thelial cells into the cleft of the comma-shaped bodies to a single system. The migration and insertion of the ureteric
form rudimentary glomerular capillary tufts (reviewed by bud into the bladder depend on Ret gene activity and Ret
Burrow, 2000). Wt1 and Pod1 may have important functions gene expression and is mediated by the action of the retinoic
in the regulation of gene transcription necessary for the dif- acid and Gata3 gene (Schultza, 2016).
ferentiation of podocytes (Ballermann, 2005).
Fibroblast growth factors (FGF) are also important for
VESICOURETHRAL UNIT
early metanephric development, especially the receptors
Fgfr1 and Fgfr2. A loss of both these receptors leads to kid- The blind end of the hindgut caudal to the point of origin of
ney agenesis. Other signaling proteins include Six1 and Sall1. the allantois expands to form the cloaca, which is separated
Six1 is a homeobox protein essential for early kidney devel- from the outside by a thin plate of tissue (the cloacal mem-
opment. Sall1 is a transcription factor that is important for brane) lying in an ectodermal depression (the proctodeum)
the development of the metanephros. Lack of Sall1 leads to under the root of the tail. At the 4-mm stage, starting at the
renal agenesis (Krause, 2015). cephalic portion of the cloaca where the allantois and gut
meet, the cloaca progressively divides into two compartments
by the caudal growth of a crescentic fold, the urorectal fold.
ANOMALIES OF THE NEPHRIC SYSTEM
The two limbs of the fold bulge into the lumen of the cloaca
Failure of the metanephros to ascend leads to an ectopic kidney. from either side, eventually meeting and fusing. The division
An ectopic kidney may be on the proper side but low (simple of the cloaca into a ventral portion (urogenital sinus) and a
ectopy) or on the opposite side (crossed ectopy) with or with- dorsal portion (rectum) is completed during the 7th week.
out fusion. Failure to rotate during ascent causes a malrotated During the development of the urorectal septum, the cloacal
kidney. Fusion of the paired metanephric masses leads to vari- membrane undergoes a reverse rotation, so that the ectoder-
ous anomalies—most commonly a “horseshoe” kidney. mal surface is no longer directed toward the developing ante-
The ureteral bud from the mesonephric duct may bifur- rior abdominal wall but gradually is turned to face caudally
cate, causing a bifid ureter at various levels depending on and slightly posteriorly. This change facilitates the subdivision
the time of the bud’s subdivision. An accessory ureteral bud of the cloaca and is brought about mainly by development of
may develop from the mesonephric duct, thereby forming the infraumbilical portion of the anterior abdominal wall and
a duplicated ureter, usually meeting the same metaneph- regression of the tail. The mesoderm that passes around the
ric mass. Rarely, each bud has a separate metanephric mass, cloacal membrane to the caudal attachment of the umbilical
resulting in supernumerary kidneys. cord proliferates and grows, forming a surface elevation, the
If the double ureteral buds are close together on the meso- genital tubercle. Further growth of the infraumbilical part
nephric duct, they open near each other in the bladder. In of the abdominal wall progressively separates the umbilical
this case, the main ureteral bud, which is the first to appear cord from the genital tubercle. The division of the cloaca is
and the most caudal on the mesonephric ducts, reaches the completed before the cloacal membrane ruptures, and its two
bladder first. It then starts to move upward and laterally and parts therefore have separate openings. The ventral part is the
is followed later by the second accessory bud as it reaches the primitive urogenital sinus, which has the shape of an elon-
urogenital sinus. The main ureteral bud (now more cranial gated cylinder and is continuous cranially with the allantois;
on the urogenital sinus) drains the lower portion of the kid- its external opening is the urogenital ostium. The dorsal part
ney. The two ureteral buds reverse their relationship as they is the rectum, and its external opening is the anus.
move from the mesonephric duct to the urogenital sinus. Traditionally, it is believed that the urogenital sinus
This is why duplicated ureters always cross (Weigert–Meyer receives the mesonephric ducts. The caudal end of the meso-
law). If the two ureteral buds are widely separated on the nephric duct distal to the ureteral bud (the common excre-
mesonephric duct, the accessory bud appears more proximal tory duct) is progressively absorbed into the urogenital sinus.
on the mesonephric duct and therefore ends in the bladder By the 7th week, the mesonephric duct and the ureteral bud
more distal than usual, with an ectopic orifice lower than the have independent opening sites. This introduces an island of
normal one. This ectopic orifice could still be in the bladder mesodermal tissue amid the surrounding endoderm of the
close to its outlet, in the urethra, or even in the genital duct urogenital sinus. As development progresses, the opening
system (Figure 2–3). A single ureteral bud that arises more of the mesonephric duct (which will become the ejaculatory
proximal than normal on the mesonephric duct can also end duct) migrates downward and medially. The opening of the
in a similar ectopic location, although this is less common. ureteral bud (which will become the ureteral orifice) migrates
Lack of development of a ureteral bud results in a solitary upward and laterally. The absorbed mesoderm of the meso-
kidney and a hemitrigone. The ureteral bud may also develop nephric duct expands with this migration to occupy the area
or migrate into the bladder, abnormally leading to a uretero- limited by the final position of these tubes (Figure 2–3).
cele. A ureterocele is a cystic dilation of the distal intramural This will later be differentiated as the trigonal structure,

▲▲Figure 2–3. Development of the ureteral bud from the mesonephric duct and the relationship of both to the urogenital
sinus. The ureteral bud appears at the 4th week. The mesonephric duct distal to this ureteral bud is gradually absorbed
into the urogenital sinus, resulting in separate endings for the ureter and the mesonephric duct. The mesonephric tissue
that is incorporated into the urogenital sinus expands and forms the trigonal tissue.
which is the only mesodermal inclusion in the endodermal supported by more recent studies that suggest the trigone is
vesicourethral unit. formed mostly from bladder smooth muscle and less so from
More recent studies suggest an alternative path of devel- the ureters. Condensation of myoblasts in the region between
opment (reviewed by McInnes and Michaud, 2009). The the openings of the ureters and Wolffian ducts at 12 weeks of
right and left common excretory ducts appear to undergo gestation gives rise to the trigone, as a single circular muscu-
gradual programmed cell death; the elimination of the com- lar layer and the muscles from the distal ureters cross midline
mon excretory ducts brings the distal ureters into immediate to form the interureteral fold (Oswald et al, 2006).
contact with the urogenital sinus epithelium. Concurrently, The urogenital sinus can be divided into two main seg-
the ureters undergo a 180° rotation around the axis of the ments. The dividing line, the junction of the combined
mesonephric duct (also known as the Wolffian duct). The Müllerian ducts with the dorsal wall of the urogenital sinus, is
distal segment of the ureters then also undergoes apoptosis. an elevation called Müller’s tubercle, which is the most fixed
As a result, this process generates a new ureteral connection reference point in the whole structure and is discussed in a
point in the urogenital sinus region that will give rise to the subsequent section. The segments are as follows:
bladder, while the Wolffian duct remains in the region giving 1. The ventral and pelvic portion forms the bladder, part of
rise to the urethra. Further growth of the bladder and ure- the urethra in males, and the whole urethra in females.
thra moves the ureteral orifices cranially, while those to the This portion receives the ureter.
Wolffian ducts move caudally. This pattern of development is

2. The urethral, or phallic, portion receives the mesonephric The part of the urogenital sinus caudal to the opening of
and the fused Müllerian ducts. This will be part of the ure- the Müllerian duct forms the vaginal vestibule and contrib-
thra in males and forms the lower fifth of the vagina and utes to the lower fifth of the vagina in females (Figure 2–5).
the vaginal vestibule in females. In males, it forms the inframontanal part of the prostatic
During the 3rd month, the ventral part of the urogenital urethra and the membranous urethra. The penile urethra
sinus starts to expand and forms an epithelial sac whose apex is formed by the fusion of the urethral folds on the ventral
tapers into an elongated, narrowed urachus. The pelvic por- surface of the genital tubercle. In females, the urethral folds
tion remains narrow and tubular; it forms the whole urethra remain separate and form the labia minora. The glandular
in females and the supramontanal portion of the prostatic urethra in males is formed by canalization of the urethral
urethra in males. The splanchnic mesoderm surrounding the plate. The bladder originally extends up to the umbilicus,
ventral and pelvic portion of the urogenital sinus begins to where it is connected to the allantois that extends into the
differentiate into interlacing bands of smooth-muscle fibers umbilical cord. The allantois usually is obliterated at the level
and an outer fibrous connective tissue coat. By the 12th week, of the umbilicus by the 15th week. The bladder then starts to
the layers characteristic of the adult urethra and bladder are descend by the 18th week. As it descends, its apex becomes
recognizable (Figure 2–4). stretched and narrowed, and it pulls on the already obliterated
▲▲Figure 2–4. Differentiation of the urogenital sinus in males. At the 5th week, the progressively growing urorectal
septum separates the urogenital sinus from the rectum. The former receives the mesonephric duct and the ureteral
bud. It retains its tubular structure until the 12th week, when the surrounding mesenchyme starts to differentiate into
the muscle fibers around the whole structure. The prostate gland develops as multiple epithelial outgrowths just above
and below the mesonephric duct. During the 3rd month, the ventral part of the urogenital sinus expands to form the
bladder proper; the pelvic part remains narrow and tubular, forming part of the urethra. (Reproduced with permission
from Tanagho EA, Smith DR: Mechanism of urinary continence. I. Embryologic, anatomic and pathologic considerations, J Urol.
1968 Nov;100(5):640–646.)

▲▲Figure 2–5. Differentiation of the urogenital sinus and the Müllerian ducts in the female embryo. At 9 weeks, the
urogenital sinus receives the fused Müllerian ducts at Müller’s tubercle (sinovaginal node), which is solidly packed with
cells. As the urogenital sinus distal to Müller’s tubercle becomes wider and shallower (15 weeks), the urethra and fused
Müllerian duct will have separate openings. The distal part of the urogenital sinus forms the vaginal vestibule and the
lower fifth of the vagina (shaded area), and that part above Müller’s tubercle forms the urinary bladder and the entire
female urethra. The fused Müllerian ducts form the uterus and the upper four-fifths of the vagina. The hymen is formed at
the junction of the sinovaginal node and the urogenital sinus.
allantois, now called the urachus. By the 20th week, the blad- anterior lobe tubules are large and show multiple branches,
der is well separated from the umbilicus, and the stretched they gradually contract and lose most of the branches. They
urachus becomes the middle umbilical ligament. continue to shrink so that at birth, they show no lumen and
appear as small, solid embryonic epithelial outgrowths. In
contrast, the tubules of the posterior lobe are fewer in num-
PROSTATE ber yet larger, with extensive branching. These tubules, as
The prostate develops as multiple solid outgrowths of the they grow, extend posterior to the developing median and lat-
urethral epithelium both above and below the entrance of the eral lobes and form the posterior aspect of the gland, which
mesonephric duct. These simple tubular outgrowths begin to may be felt rectally.
develop in five distinct groups at the end of the 11th week and Prostate development results from complex interaction
are complete by the 16th week (112-mm stage). They branch between urogenital sinus epithelium and mesenchyme in
and rebranch, ending in a complex duct system that encoun- the presence of androgens (reviewed by Cunha et al, 2004,
ters the differentiating mesenchymal cells around this seg- and Thomson, 2008). Early in development, the androgen
ment of the urogenital sinus. These mesenchymal cells start receptors are solely expressed in the urogenital sinus mesen-
to develop around the tubules by the 16th week and become chyme. Under the influence of androgen, the mesenchyme
denser at the periphery to form the prostatic capsule. By the induces epithelial bud formation, regulates the growth and
22nd week, the muscular stroma is considerably developed, branching of epithelial bud, promotes differentiation of a
and it continues to increase progressively until birth. secretory epithelium, and specifies differential expression
From the five groups of epithelial buds, five lobes are eventu- of prostatic secretory proteins. Genome-wide analyses have
ally formed: anterior, posterior, median, and two lateral lobes. revealed critical molecular events in prostate development.
Initially, these lobes are widely separated, but later they meet, These include Nkx3.1, Sox, and homeobox genes for ductal
with no definite septa dividing them. Tubules of each lobe do morphology development; sonic hedgehog, fibroblast growth
not intermingle with each other but simply lie side by side. factor and the Wnt5a gene for bud development; and bone
The anterior lobe tubules begin to develop simultaneously morphogenic protein and notch genes for branching (Meeks,
with those of the other lobes. In the early stages, although the 2011). Other factors such as activin A serve to inhibit ductal

branching in order to maintain tissue homeostasis and regu- mesonephros is converted into a gonadal mesentery known
lated growth in the prostate. as the mesorchium. The cells of the germinal epithelium
grow into the underlying mesenchyme and form cordlike
ANOMALIES OF THE VESICOURETHRAL UNIT masses. These are radially arranged and converge toward the
mesorchium, where a dense portion of the blastemal mass
Failure of the cloaca to subdivide is rare and results in a
is also emerging as the primordium of the rete testis. A net-
persistent cloaca. Incomplete subdivision is more frequent,
work of strands soon forms that is continuous with the testis
ending with rectovesical, rectourethral, or rectovestibular
cords. The latter also split into three to four daughter cords.
fistulas (usually with imperforate anus or anal atresia).
These eventually become differentiated into the seminifer-
Failure of descent or incomplete descent of the blad-
ous tubules by which the spermatozoa are produced. The rete
der leads to a urinary umbilical fistula (urachal fistula),
testis unites with the mesonephric components that will form
urachal cyst, or urachal diverticulum depending on the
the male genital ducts, as discussed in a subsequent section
stage and degree of maldescent.
(Figure 2–6).
Development of the genital primordia in an area more
If the gonad develops into an ovary, it (like the testis) gains
caudal than normal can result in formation of the corpora
a mesentery (mesovarium) and settles in a more caudal posi-
cavernosa just caudal to the urogenital sinus outlet, with the
tion. During the 9th week the internal blastema differentiates
urethral groove on its dorsal surface. This defect results in
in the into a primary cortex beneath the germinal epithelium
complete or incomplete epispadias depending on its degree.
and a loose primary medulla. A compact cellular mass bulges
A more extensive defect results in bladder exstrophy. Failure
from the medulla into the mesovarium and establishes the
of fusion of urethral folds leads to various grades of hypospa-
primitive rete ovarii. At 3–4 months of age, the internal cell
dias. This defect, because of its mechanism, never extends
mass becomes young ova. A new definitive cortex is formed
proximal to the bulbous urethra. This is in contrast to epi-
from the germinal epithelium as well as from the blastema
spadias, which usually involves the entire urethra up to the
in the form of distinct cellular cords (Pflüger’s tubes), and a
internal meatus.
permanent medulla is formed. The cortex differentiates into
ovarian follicles containing ova.
GONADS
Genetically, in the presence of a Y chromosome, SRY (as
Most of the structures that make up the embryonic genital sys- known as testis determining factor [TDF]) induces the upreg-
tem have been taken over from other systems, and their read- ulation of Sox9 in the undifferentiated gonad (reviewed by
aptation to genital function is a secondary and relatively late Sekido, 2010). This in turn upregulates the expression of FGF9
phase in their development. The early differentiation of such and increases PGD2 synthesis, both helping to maintain Sox9
structures is therefore independent of sexuality. Furthermore, expression. Sox9 directs the differentiation of cells into Sertoli
each embryo is at first morphologically bisexual, possessing all cell by activating several downstream genes such as Amh,
the necessary structures for either sex. The development of one Cbln4, FGF9, and Ptgds. In the absence of a Y chromosome
set of sex primordia and the gradual involution of the other are and SRY, Rspo1 is upregulated in the undifferentiated gonads
determined by the sex of the gonad. (reviewed by Nef and Vassalli, 2009). Rspo1 is required for
The sexually undifferentiated gonad is a composite struc- Wnt4 expression, and together they activate β-catenin, which,
ture. Male and female potentials are represented by specific in turn, suppresses the formation of the testis cords by inhibit-
histologic elements (medulla and cortex) that have alterna- ing Sox9 and FGF9. A second pathway involving the upregu-
tive roles in gonadogenesis. Normal differentiation involves lation of Foxl2 also serves to inhibit Sox 9 and FGF9 activity,
the gradual predominance of one component. contributing to the development of the ovary.
The primitive sex glands appear during the 5th and 6th New genetic data on human sex determination from
weeks within a localized region of the thickening known as modern gene sequencing will create opportunities for the
the urogenital ridge (this contains both the nephric and the development of mechanistic models and should lead to
genital primordia). At the 6th week, the gonad consists of better understanding of this complex process (reviewed by
a superficial germinal epithelium and an internal blastema. Bashamboo, 2017).
The blastemal mass is derived mainly from proliferative
ingrowth from the superficial epithelium, which comes loose ▶▶Descent of the Gonads
from its basement membrane.
A. Testis
During the 7th week, the gonad begins to assume the
characteristics of a testis or ovary. Differentiation of the In addition to its early caudal migration, the testis later leaves
ovary usually occurs somewhat later than differentiation of the abdominal cavity and descends into the scrotum. By the
the testis. 3rd month of fetal life, the testis is located retroperitoneally
If the gonad develops into a testis, the gland increases in in the false pelvis. A fibromuscular band (the gubernacu-
size and shortens into a more compact organ while achiev- lum) extends from the lower pole of the testis through the
ing a more caudal location. Its broad attachment to the developing muscular layers of the anterior abdominal wall

▲▲Figure 2–6. Transformation of the undifferentiated genital system into the definitive male and female systems.

to terminate in the subcutaneous tissue of the scrotal swell- the 4-mm stage) joins the ventral part of the cloaca, which
ing. The gubernaculum also has several other subsidiary will be the urogenital sinus. This duct gives rise to the ureteral
strands that extend to adjacent regions. Just below the bud close to its caudal end. The ureteral bud grows cranially
lower pole of the testis, the peritoneum herniates as a and meets metanephrogenic tissue. The part of each meso-
diverticulum along the anterior aspect of the gubernacu- nephric duct caudal to the origin of the ureteric bud becomes
lum, eventually reaching the scrotal sac through the ante- absorbed into the wall of the primitive urogenital sinus so
rior abdominal muscles (the processus vaginalis). The testis that the mesonephric duct and ureter open independently.
remains at the abdominal end of the inguinal canal until the This is achieved at the 15-mm stage (7th week). During this
7th month. It then passes through the inguinal canal behind period, starting at the 10-mm stage, the Müllerian ducts start
(but invaginating) the processus vaginalis. Normally, it to develop. They reach the urogenital sinus relatively late—at
reaches the scrotal sac by the end of the 8th month. the 30-mm stage (9th week); their partially fused blind ends
producing the elevation called Müller’s tubercle. Müller’s
B. Ovary tubercle is the most constant and reliable point of reference
in the whole system.
In addition to undergoing an early internal descent, the ovary If the gonad starts to develop into a testis (17-mm stage,
becomes attached through the gubernaculum to the tissues 7th week), the Wolffian duct will start to differentiate into
of the genital fold and then attaches itself to the developing the male duct system, forming the epididymis, vas deferens,
uterovaginal canal at its junction with the uterine (fallopian) seminal vesicles, and ejaculatory ducts, when the Müllerian
tubes. This part of the gubernaculum between the ovary and duct proceeds toward its junction with the urogenital sinus
uterus becomes the ovarian ligament; the part between the and immediately starts to degenerate. Only its upper and
uterus and the labia majora becomes the round ligament lower ends persist, the former as the appendix testis and the
of the uterus. These ligaments prevent extra-abdominal latter as part of the prostatic utricle.
descent, and the ovary enters the true pelvis. It eventually lies If the gonad starts to differentiate into an ovary (22-mm
posterior to the uterine tubes on the superior surface of the stage, 8th week), the Müllerian duct system forms the uter-
urogenital mesentery, which has descended with the ovary ine (fallopian) tubes, uterus, and most of the vagina. The
and now forms the broad ligament. A small processus vag- Wolffian ducts, aside from their contribution to the urogeni-
inalis forms and passes toward the labial swelling, but it is tal sinus, remain rudimentary.
usually obliterated at full term.
MALE DUCT SYSTEM
GONADAL ANOMALIES
The gonad may either (1) not develop (agenesis), (2) develop ▶▶Epididymis
incompletely (hypogenesis), (3) develop incorrectly (dysgen- Because of the proximity of the differentiating gonads and the
esis), or (4) develop correctly, but then suffer an embryologic nephric duct, some of the mesonephric tubules are retained
injury (eg, intrauterine torsion). Supernumerary gonads are as the efferent ductules, and their lumens become continu-
rare. The commonest anomaly involves descent of the gonads, ous with those of the rete testis. These tubules, together with
especially the testis. Retention of the testis in the abdomen or the part of the mesonephric duct into which they empty, will
arrest of its descent at any point along its natural pathway form the epididymis. Each coiled ductule makes a conical
is called cryptorchidism, which may be either unilateral or mass known as the lobule of the epididymis. The cranial end
bilateral. If the testis does not follow the main gubernacular of the mesonephric duct becomes highly convoluted, com-
structure but follows one of its subsidiary strands, it will end pleting the formation of the epididymis. This is an example of
in an abnormal position, resulting in an ectopic testis. direct inclusion of a nephric structure into the genital system.
Failure of union between the rete testis and mesonephros Additional mesonephric tubules, both cephalad and caudal
results in a testis separate from the male genital ducts (the to those that were included in the formation of the epididy-
epididymis) and azoospermia. mis, remain as rudimentary structures, that is, the appendix
of the epididymis and the paradidymis.
GENITAL DUCT SYSTEM
Alongside the indifferent gonads, there are, early in embry- ▶▶Vas Deferens, Seminal Vesicles, and
Ejaculatory Ducts
onic life, two different yet closely related ducts. One is pri-
marily a nephric duct (Wolffian duct), yet it also serves as a The mesonephric duct caudal to the portion forming the
genital duct if the embryo develops into a male. The other epididymis forms the vas deferens. Shortly before this duct
(Müllerian duct) is primarily a genital structure from the joins the urethra (urogenital sinus), a localized dilatation
start. (ampulla) develops, and the saccular convoluted structure
Both ducts grow caudally to join the primitive urogenital that will form the seminal vesicle is evaginated from its wall.
sinus. The Wolffian duct (known as the pronephric duct at The mesonephric duct between the origin of the seminal

vesicle and the urethra forms the ejaculatory duct. The whole tubes (fallopian tubes, oviducts) are the cephalic two-thirds
mesonephric duct now achieves its characteristic thick of the Müllerian ducts (Figure 2–6).
investment of smooth muscle, with a narrow lumen along
most of its length. ANOMALIES OF THE GONADAL DUCT SYSTEM
Both above and below the point of entrance of the meso-
Nonunion of the rete testis and the efferent ductules can
nephric duct into the urethra, multiple outgrowths of ure-
occur and, if bilateral, causes azoospermia and sterility.
thral epithelium mark the beginning of the development of
Failure of the Müllerian ducts to approximate or to fuse com-
the prostate. As these epithelial buds grow, they meet the
pletely can lead to various degrees of duplication in the geni-
developing muscular fibers around the urogenital sinus,
tal ducts. Congenital absence of one or both uterine tubes or
and some of these fibers become entangled in the branch-
of the uterus or vagina occurs rarely.
ing tubules of the growing prostate and become incorporated
Arrested development of the infratubercular segment of
into it, forming its muscular stroma (Figure 2–4).
the urogenital sinus leads to its persistence, with the urethra
and vagina having a common duct to the outside (urogenital
FEMALE DUCT SYSTEM sinus).
The Müllerian ducts, which are a paired system, are seen
alongside the mesonephric duct. It is not known whether EXTERNAL GENITALIA
they arise directly from the mesonephric ducts or separately During the 8th week, external sexual differentiation begins
as an invagination of the celomic epithelium into the paren- to occur. Not until 3 months, however, do the progressively
chyma lateral to the cranial extremity of the mesonephric developing external genitalia attain characteristics that can
duct, but the latter theory is favored. The Müllerian duct be recognized as distinctively male or female. During the
develops and runs lateral to the mesonephric duct. Its open- indifferent stage of sexual development, three small protu-
ing into the celomic cavity persists as the peritoneal ostium berances appear on the external aspect of the cloacal mem-
of the uterine tube (later it develops fimbriae). The other brane. In front is the genital tubercle, and on either side of the
end grows caudally as a solid tip and then crosses in front of membrane are the genital swellings.
the mesonephric duct at the caudal extremity of the meso- With the breakdown of the urogenital membrane (17-mm
nephros. It continues its growth in a caudomedial direction stage, 7th week), the primitive urogenital sinus achieves a
until it meets and fuses with the Müllerian duct of the oppo- separate opening on the undersurface of the genital tubercle.
site side. The fusion is partial at first, so there is a tempo-
rary septum between the two lumens. This later disappears,
leaving one cavity that will form the uterovaginal canal. The
MALE EXTERNAL GENITALIA
potential lumen of the vaginal canal is completely packed The urogenital sinus opening extends on the ventral aspect
with cells. The solid tip of this cord pushes the epithelium of the genital tubercle as the urethral groove. The primitive
of the urogenital sinus outward, where it becomes Müller’s urogenital orifice and the urethral groove are bound on either
tubercle (33-mm stage, 9th week). The Müllerian ducts fuse side by the urethral folds. The genital tubercle becomes elon-
at the 63-mm stage (13th week), forming the sinovaginal gated to form the phallus. The corpora cavernosa is indicated
node, which receives a limited contribution from the uro- in the 7th week as paired mesenchymal columns within the
genital sinus (this contribution forms the lower fifth of the shaft of the penis. By the 10th week, the urethral folds start
vagina). to fuse from the urogenital sinus orifice toward the tip of the
The urogenital sinus distal to Müller’s tubercle, originally phallus. At the 14th week, the fusion is complete and results
narrow and deep, shortens, widens, and opens to form the in the formation of the penile urethra. The corpus spongio-
floor of the pudendal or vulval cleft. This results in separate sum results from the differentiation of the mesenchymal
openings for the vagina and urethra and brings the vaginal masses around the formed penile urethra.
orifice to its final position nearer the surface. At the same The glans penis becomes defined by the development of a
time, the vaginal segment increases appreciably in length. circular coronary sulcus around the distal part of the phallus.
The vaginal vestibule is derived from the infratubercular seg- The urethral groove and the fusing folds do not extend
ment of the urogenital sinus (in males, the same segment will beyond the coronary sulcus. The glandular urethra develops
form the inframontanal part of the prostatic urethra and the as a result of canalization of an ectodermal epithelial cord that
membranous urethra). The labia minora are formed from the has grown through the glans. This canalization reaches and
urethral folds (in males they form the pendulous urethra). communicates with the distal end of the previously formed
The hymen is the remnant of the Müllerian tubercle. The penile urethra. During the 3rd month, a fold of skin at the
lower fifth of the vagina is derived from the portion of the base of the glans begins growing distally and, 2 months later,
urogenital sinus that combines with the sinovaginal node. surrounds the glans. This forms the prepuce. Meanwhile, the
The remainder of the vagina and the uterus are formed from genital swellings shift caudally and are recognizable as scrotal
the lower (fused) third of the Müllerian ducts. The uterine swellings. They meet and fuse, resulting in the formation of

the scrotum, with two compartments partially separated by Michos O: Kidney development: From ureteric bud formation
a median septum and a median raphe, indicating their line to branching morphogenesis. Curr Opin Genet Dev 2009;19:
of fusion. 484–490.
Mcinnes RR, Michaud JL: Plumbing in the embryo: Development
defects of the urinary tracts. Clin Genet 2009;75:307–317.
FEMALE EXTERNAL GENITALIA
Nef S, Vassalli JD: Complementary pathways in mammalian female
Until the 8th week, the appearance of the female external geni- sex determination. J Biol 2009;8:74.
talia closely resembles that of the male genitalia except that the Nyengaard JR, Bendtsen TF: Glomerular number and size in relation
urethral groove is shorter. The genital tubercle, which becomes to age, kidney weight, and body surface in normal man. Anat Rec
bent caudally and lags in development, becomes the clitoris. As 1992;232(2):194–201.
in males (albeit on a minor scale), mesenchymal columns differ- Oswald J et al: Reevaluation of the fetal muscle development of the
entiate into corpora cavernosa, and a coronary sulcus identifies vesical trigone. J Urol 2006;176(3):1166–1170.
the glans clitoridis. The most caudal part of the urogenital sinus Reddy PP, Mandell J: Prenatal diagnosis: Therapeutic implications.
Urol Clin North Am 1998;25:171.
shortens and widens, forming the vaginal vestibule. The urethral
folds do not fuse but remain separate as the labia minora. The Sekido R: SRY: A transcriptional activator of mammalian testis deter-
mination. Int J Biochem Cell Biol 2010;42(3):417–420.
genital swellings meet in front of the anus, forming the posterior
Shah MM et al: Branching morphogenesis and kidney disease.
commissure, while the swellings enlarge and remain separated Development 2004;131(7):1449–1462.
on either side of the vestibule and form the labia majora.
Stephens FD: Embryopathy of malformations. J Urol 1982;127:13.
Stephens FD: Congenital Malformations of the Urinary Tract. Prae-
ANOMALIES OF THE EXTERNAL GENITALIA ger, New York, 1983.
Absence or duplication of the penis or clitoris is very rare. Tanagho EA: Embryologic development of the urinary tract. In: Ball
More commonly, the penis remains rudimentary or the cli- TP (ed): AUA Update Series. American Urological Association,
Philadelphia, 1982.
toris shows hypertrophy. These anomalies may be seen alone
Tanagho EA: Developmental anatomy and urogenital abnormalities.
or, more frequently, in association with differences of sex
In: Raz S (ed): Female Urology. 2nd ed. Saunders, Philadelphia,
development (DSD). Concealed penis and transposition of 1986.
penis and scrotum are relatively rare anomalies. Thomson AA: Mesenchymal mechanisms in prostate organogenesis.
Failure or incomplete fusion of the urethral folds results Differentiation 2008;76(6):587–598.
in hypospadias (see preceding discussion). Penile develop- Vaughan ED Jr, Middleton GW: Pertinent genitourinary embryology:
ment is also anomalous in cases of epispadias and exstrophy Review for practicing urologist. Urology 1975;6:139.
(see preceding discussion).
Anomalies of the Nephric System

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31
3
Symptoms of Disorders of
the Genitourinary Tract
In the workup of any patient, the history is of paramount General malaise may be noted with tumors, chronic
importance; this is particularly true in urology. It is neces- pyelonephritis, or renal failure. The presence of many of
sary to discuss here only those urologic symptoms that are these symptoms may be compatible with human immunode-
apt to be brought to the physician’s attention by the patient. ficiency virus (HIV; see Chapter 17).
It is important to know not only whether the disease is acute
or chronic but also whether it is recurrent, since recurring LOCAL AND REFERRED PAIN
symptoms may represent acute exacerbations of chronic
disease. Two types of pain have their origins in the genitourinary
Obtaining the history is an art that depends on the skill organs: local and referred. The latter is especially common.
and methods used to elicit information. The history is only Local pain is felt in or near the involved organ. Thus, the
as accurate as the patient’s ability to describe the symptoms. pain from a diseased kidney (T10–T12, L1) is felt in the cos-
This subjective information is important in establishing an tovertebral angle and in the flank in the region of and below
accurate diagnosis. the 12th rib. Pain from an inflamed testicle is felt in the gonad
itself.
SYSTEMIC MANIFESTATIONS Referred pain originates in a diseased organ but is felt at
some distance from that organ. The ureteral colic (Figure 3–1)
Symptoms of fever and weight loss should be sought. The
caused by a stone in the upper ureter may be associated with
presence of fever associated with other symptoms of urinary
severe pain in the ipsilateral testicle; this is explained by the
tract infection may be helpful in evaluating the site of the
common innervation of these two structures (T11–T12).
infection. Simple acute cystitis is essentially an afebrile dis-
A stone in the lower ureter may cause pain referred to the
ease. Acute pyelonephritis or prostatitis is apt to cause high
scrotal wall; in this instance, the testis itself is not hyperes-
temperatures (≤40°C [104°F]), often accompanied by violent
thetic. The burning pain with voiding that accompanies
chills. Infants and children who have acute pyelonephritis
acute cystitis is felt in the distal urethra in females and in the
may have high temperatures without other localizing symp-
glandular urethra in males (S2–S3).
toms or signs. Such a clinical picture, therefore, invariably
Abnormalities of a urologic organ can also cause pain in
requires bacteriologic study of the urine.
any other organ (eg, gastrointestinal, gynecologic) that has a
A history of unexplained attacks of fever occurring even
sensory nerve supply common to both (Figures 3–2 and 3–3).
years before may otherwise represent asymptomatic pyelo-
nephritis. Renal carcinoma sometimes causes fever that may
reach 39°C (102.2°F) or more. The absence of fever does not ▶▶Kidney Pain (Figure 3–1)
by any means rule out renal infection, for it is the rule that Typical renal pain is felt as a dull and constant ache in the
chronic pyelonephritis does not cause fever. costovertebral angle just lateral to the sacrospinalis muscle
Weight loss is to be expected in the advanced stages of and just below the 12th rib. This pain often spreads along
cancer, but it may also be noticed when renal insufficiency the subcostal area toward the umbilicus or lower abdominal
due to obstruction or infection supervenes. In children who quadrant. It may be expected in the renal diseases that cause
have “failure to thrive” (low weight and less than average sudden distention of the renal capsule. Acute pyelonephritis
height for their age), chronic obstruction, urinary tract infec- (with its sudden edema) and acute ureteral obstruction (with
tion, or both should be suspected. its sudden renal back pressure) both cause this typical pain.

▲▲Figure 3–1. Referred pain from kidney (dotted areas) and ureter (shaded areas).
It should be pointed out, however, that many urologic renal The physician may be able to judge the position of a ure-
diseases are painless because their progression is so slow that teral stone by the history of pain and the site of referral. If the
sudden capsular distention does not occur. Such diseases stone is lodged in the upper ureter, the pain radiates to
include cancer, chronic pyelonephritis, staghorn calculus, the testicle, since the nerve supply of this organ is simi-
tuberculosis, polycystic kidney, and hydronephrosis due to lar to those of the kidney and upper ureter (T11–T12).
chronic ureteral obstruction. With stones in the midportion of the ureter on the right side,
the pain is referred to McBurney’s point and may therefore
▶▶Ureteral Pain (Figure 3–1) simulate appendicitis; on the left side, it may resemble diver-
ticulitis or other diseases of the descending or sigmoid colon
Ureteral pain is typically stimulated by acute obstruction (pas-
(T12, L1). As the stone approaches the bladder, inflamma-
sage of a stone or a blood clot). In this instance, there is back
tion and edema of the ureteral orifice ensue, and symptoms
pain from renal capsular distention combined with severe col-
of vesical irritability such as urinary frequency and urgency
icky pain (due to renal pelvic and ureteral muscle spasm) that
may occur. It is important to realize, however, that in mild
radiates from the costovertebral angle down toward the lower
ureteral obstruction, as seen in the congenital stenoses, there
anterior abdominal quadrant, along the course of the ureter.
is usually no pain, either renal or ureteral.
In men, it may also be felt in the bladder, scrotum, or testicle.
In women, it may radiate into the vulva. The severity and col-
icky nature of this pain are caused by the hyperperistalsis and ▶▶Vesical Pain
spasm of this smooth-muscle organ as it attempts to rid itself The overdistended bladder of the patient in acute urinary
of a foreign body or to overcome obstruction. retention causes agonizing pain in the suprapubic area. Other

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corresponding to those of the maxilla can be clearly traced in the
labium.
Fig. 52.—Maxilla and lower lip of Coleoptera. A, Maxilla of Passalus: a,

cardo; b, stipes; c, palpiger; d, palpus; e, inner or inferior lobe or
lacinia; f, outer or superior lobe or galea: B, Labium of Harpalus
caliginosus: a, mentum; b, hypoglottis; c, palpiger (support of the
labial palp); d, palp; e, ligula; f, paraglossa.
The mentum is an undivided, frequently very hard, piece, continuous

with either the submentum or the gula, and anterior to this are placed
the other parts, viz. the labial palpi and their supports, the palpigers;
beyond and between these exists a central piece (Fig. 52, B, e),
about whose name some difference of opinion prevails, but which
may be called the ligula (languette of French authors), and on each
side of this is a paraglossa. In the Orthoptera the single median
piece—the ligula of Coleopterists—is represented by two divided
parts. In some Insects (many Coleoptera) there is interposed
between the mentum and the palpigers a piece called the hypoglottis
(Fig. 52, B, b). It is not so well ascertained as it should be, that the
pieces of the lower lip bearing the same names in different Orders
are in all cases really homologous, and comparison suggests that
the hypoglottis of Coleoptera may possibly represent the piece
corresponding to the mentum of Orthopterists, the so-called mentum
of beetles being in that case the submentum of Orthopterists.
There is another part of the mouth to which we may call special

attention, as it has recently attracted more attention than it formerly
did; it is a membranous lobe in the interior of the mouth, very
conspicuous in Orthoptera, and called the tongue, lingua, or
hypopharynx; it reposes, in the interior of the mouth (Fig. 51, o), on
the middle parts of the front of the labium; it is probably not entirely
lost in Coleoptera, but enters into the composition of the complex
middle part of the lip by amalgamation with the paraglossae. It has
recently been proposed to treat this lingua as the morphological
equivalent of the labium or of the maxillae, giving it the name of the
endolabium, but the propriety of this course remains to be proved;[20]
the view is apparently suggested chiefly by the structure of the
mouth of Hemimerus, a very rare and most peculiar Insect that has
not as yet been sufficiently studied.
As the maxillae and labium are largely used by taxonomists in the

systematic arrangement of the mandibulate Insects, we give a figure
of them as seen in Coleoptera, where the parts, though closely
amalgamated, can nevertheless be distinguished. This Fig. 52
should be compared with Fig. 51.
In speaking of the segments of the body we pointed out that they

were not separate parts but constituted an uninterrupted whole, and
it is well to remark here that this is also true of the gnathites.
Although the mouth parts are spoken of as separate pieces, they
really form only projections from the great body wall. Fig. 51, B,
shows the intimate connexion that exists between the maxillae and
labium; the continuity of the mandibles with the membrane of the
buccal cavity is capable of very easy demonstration.
The head bears, besides the pieces we have considered, a pair of

antennae. These organs, though varying excessively in form, are
always present in the adult Insect, and exist even in the majority of
young Insects. They are very mobile, highly sensitive organs, situate
on or near the front part of the head. The antennae arise in the
embryo from the procephalic lobes, the morphological import of
which parts is one of the most difficult points connected with Insect
embryology.
The eyes of Insects are of two sorts, simple and compound. The
simple eyes, or ocelli, vary in number from one to as many as
eighteen or twenty; when thus numerous they are situated in groups
on each side of the head. In their most perfect form, as found in adult
aculeate Hymenoptera, in Orthoptera and Diptera, ocelli are usually
two or three in number, and present the appearance of small,
perfectly transparent lenses inserted in the integument. In their
simplest form they are said to consist of some masses of pigment in
connexion with a nerve.
Fig. 53.—Two ommatidia from the eye of Colymbetes fuscus, × 160.

(After Exner.) a, Cornea; b, crystalline cone; c, rhabdom; d,
fenestrate membrane with nerve structures below it; e, iris-
pigment; f, retina-pigment.
The compound, or facetted, eyes are the most remarkable of all the
structures of the Insect, and in the higher and more active forms,
such as the Dragon-flies and hovering Diptera, attain a complexity
and delicacy of organisation that elicit the highest admiration from
every one who studies them. They are totally different in structure
and very distinct in function from the eyes of Vertebrata, and are
seated on very large special lobes of the brain (see Fig. 65), which
indeed are so large and so complex in structure that Insects may be
described as possessing special ocular brains brought into relation
with the lights, shades, and movements of the external world by a
remarkably complex optical apparatus. This instrumental part of the
eye is called the dioptric part in contradistinction from the percipient
portion, and consists of an outer corneal lens (a, Fig. 53), whose
exposed surface forms one of the facets of the eye; under the lens is
placed the crystalline cone (b), this latter being borne on a rod-like
object (c), called the rhabdom. There are two layers of pigment, the
outer (e), called the iris-pigment, the inner (f), the retinal-pigment;
underneath, or rather we should say more central than, the
rhabdoms is the fenestrate membrane (d), beyond which there is an
extremely complex mass of nerve-fibres; nerves also penetrate the
fenestrate membrane, and their distal extremities are connected with
the delicate sheaths by one of which each rhabdom is surrounded,
the combination of sheath and nerves forming a retinula. Each set of
the parts above the fenestrate membrane constitutes an
ommatidium, and there may be many of these ommatidia in an eye;
indeed, it is said that the eye of a small beetle, Mordella, contains as
many as 25,000 ommatidia. As a rule the larvae of Insects with a
complete metamorphosis bear only simple eyes. In the young of
Dragon-flies, as well as of some other Insects having a less perfect
metamorphosis, the compound eyes exist in the early stages, but
they have then an obscure appearance, and are probably
functionally imperfect.
In the interior of the head there exists a horny framework called the
tentorium, whose chief office apparently is to protect the brain. It is
different in kind according to the species. The head shows a
remarkable and unique relation to the following segments. It is the
rule in Insect structure that the back of a segment overlaps the front
part of the one following it; in other words, each segment receives
within it the front of the one behind it. Though this is one of the most
constant features of Insect anatomy, it is departed from in the case
of the head, which may be either received into, or overlapped by, the
segment following it, but never itself overlaps the latter. There is
perhaps but a single Insect (Hypocephalus, an anomalous beetle) in
which the relation between the head and thorax can be considered
to be at all similar to that which exists between each of the other
segments of the body and that following it; and even in
Hypocephalus it is only the posterior angles of the head that overlap
the thorax. Although the head usually appears to be very closely
connected with the thorax, and is very frequently in repose received
to a considerable extent within the latter, it nevertheless enjoys great
freedom of motion; this is obtained by means of a large membrane,
capable of much corrugation, and in which there are seated some
sclerites, so arranged as to fold together and occupy little space
when the head is retracted, but which help to prop and support it
when extended for feeding or other purposes. These pieces are
called the cervical sclerites or plates. They are very largely
developed in Hymenoptera, in many Coleoptera, and in Blattidæ,
and have not yet received from anatomists a sufficient amount of
attention. Huxley suggested that they may be portions of head
segments.
Fig. 54.—Extended head and front of thorax of a beetle, Euchroma: a,

back of head; b, front of pronotum; c, chitinous retractile band; d,
cervical sclerites.
Thorax.
The thorax, being composed of the three consecutive rings behind

the head, falls naturally into three divisions—pro-, meso-, and
metathorax. These three segments differ greatly in their relative
proportions in different Insects, and in different stages of the same
Insect's life. In their more highly developed conditions each of the
three divisions is of complex structure, and the sclerites of which it is
externally made up are sufficiently constant in their numbers and
relative positions to permit of their identification in a vast number of
cases; hence the sclerites have received names, and their
nomenclature is of practical importance, because some, if not all, of
these parts are made use of in the classification of Insects. Each
division of the thorax has an upper region, called synonymically
dorsum, notum, or tergum; an inferior or ventral region, called
sternum; and on each side a lateral region, the pleuron. These
regions of each of the three thoracic divisions are further
distinguished by joining to their name an indication of the segment
spoken of, in the form of the prefixes pro-, meso-, and meta-; thus
the pronotum, prosternum, and propleura make up the prothorax.
The thoracic regions are each made up of sclerites whose
nomenclature is due to Audouin.[21] He considered that every
thoracic ring is composed of the pieces shown in Fig. 55, viz. (1) the
sternum (B', a), an unpaired ventral piece; (2) the notum (A),
composed of four pieces placed in consecutive longitudinal order
(A'), and named praescutum (a), scutum (b), scutellum (c), and post-
scutellum (d); (3) lateral pieces, of which he distinguished on each
side an episternum (B', c), epimeron (e), and parapteron (d), these
together forming the pleuron. We give Audouin's Figure, but we
cannot enter on a full discussion of his views as to the thorax; they
have become widely known, though the constancy of the parts is not
so great as he supposed it would prove to be. Sometimes it is
impossible to find all the elements he thought should be present in a
thoracic ring, while in other cases too many sclerites exist. As a rule
the notum of the meso- and metathoraces is in greater part
composed of two pieces, the scutum and the scutellum; while in the
pronotum only one dorsal piece can be satisfactorily distinguished,
though a study of the development may show that really two are
frequently, if not usually, present. On the other hand, one, or more, of
the notal sclerites in some cases shows evidence of longitudinal
division along the middle. The sternum or ventral piece, though
varying greatly in form, is the most constant element of a thoracic
segment, but it has sometimes the appearance of consisting of two
parts, an anterior and a posterior. The pleuron nearly always
consists quite evidently of two parts, the episternum, the more
anterior and inferior, and the epimeron.[22] The relations between
these two parts vary much; in some cases the episternum is
conspicuously the more anterior, while in others the epimeron is
placed much above it, and may extend nearly as far forwards as it. It
may be said, as a rule, that when the sternum extends farther
backwards than the notum, the epimeron is above the episternum,
as in many Coleoptera; but if the sternum be anterior to the notum,
then the episternum is superior to the epimeron, as in dragon-flies.
We would here again reiterate the fact that these "pieces" are really
not separate parts, but are more or less indurated portions of a
continuous integument, which is frequently entirely occupied by
them; hence a portion of a sclerite that in one species is hard, may in
an allied form be wholly or partly membranous, and in such case its
delimitation may be very evident on some of its sides, and quite
obscure on another.
Fig. 55.—Mesothorax of Dytiscus, after Audouin. A, notum; A', pieces

of the notum separated: a, praescutum; b, scutum; c, scutellum; d,
post-scutellum: B, the sternum and pleura united; B', their parts
separated: a, sternum; c, episternum; d, parapteron; e, epimeron.
The parapteron of Audouin does not appear to be really a distinct

portion of the pleuron; in the case of Dytiscus it is apparently merely
a thickening of an edge. Audouin supposed this part to be specially
connected with the wing-articulation, and the term has been
subsequently used by other writers in connexion with several little
pieces that exist in the pleural region of winged Insects.
The prothorax is even more subject to variation in its development

than the other divisions of the thorax are. In the Hymenoptera the
prosternum is disconnected from the pronotum and is capable,
together with the first pair of legs, of movement independent of its
corresponding dorsal part, the pronotum, which in this Order is
always more or less completely united with the meso-thorax; in the
Diptera the rule is that the three thoracic segments are closely
consolidated into one mass. In the majority of Insects the prothorax
is comparatively free, that is to say, it is not so closely united with the
other two thoracic segments as they are with one another. The three
thoracic rings are seen in a comparatively uniform state of
development in a great number of larvae; also in the adult stages of
some Aptera, and among winged insects in some Neuroptera such
as the Embiidae, Termitidae, and Perlidae. In Lepidoptera the
pronotum bears a pair of erectile processes called patagia; though
frequently of moderately large size, they escape observation, being
covered with scales and usually closely adpressed to the sides of the
pronotum.
The two great divisions of the body—the mesothorax and the

metathorax—are usually very intimately combined in winged Insects,
and even when the prothorax is free, as in Coleoptera, these
posterior two thoracic rings are very greatly amalgamated. In the
higher forms of the Order just mentioned the mesosternum and
mesopleuron become changed in direction, and form as it were a
diaphragm closing the front of the metasternum. The meso- and
meta-thorax frequently each bear a pair of wings.
We have described briefly and figured (Fig. 55) the sclerites of the
mesothorax, and those of the metathorax correspond fairly well with
them. In addition to the sclerites usually described as constituting
these two thoracic divisions, there are some small pieces at the
bases of the wings. Jurine discriminated and named no less than
seven of these at the base of the anterior wing of a Hymenopteron.
One of them becomes of considerable size and importance in the
Order just mentioned, and seems to be articulated so as to exert
pressure on the base of the costa of the wing. This structure attains
its maximum of development in a genus (? nondescript) of Scoliidae,
as shown in Fig. 56. The best name for this sclerite seems to be that
proposed by Kirby and Spence, tegula. Some writers call it
paraptère, hypoptère, or squamule, and others have termed it
patagium; this latter name is, however, inadmissible, as it is applied
to a process of the prothorax we have already alluded to.
Fig. 56.—Head and thorax of wasp from Bogota: t, tegula; b, base of
wing.
To complete our account of the structure of the thorax it is necessary

to mention certain hard parts projecting into its interior, but of which
there is usually little or no trace externally. A large process in many
Insects projects upwards from the sternum in a forked manner. It
was called by Audouin the entothorax; some modern authors prefer
the term apophysis. Longitudinal partitions of very large size,
descending from the dorsum into the interior, also exist; these are
called phragmas, and are of great importance in some Insects with
perfect flight, such as Hymenoptera, Lepidoptera, and Diptera. There
is no phragma in connection with the pronotum, but behind this part
there may be three. A phragma has the appearance of being a fold
of the dorsum; it serves as an attachment for muscles, and may
probably be of service in other ways. More insignificant projections
into the interior are the little pieces called apodemes (Fig. 57, e);
these are placed at the sides of the thorax near the wings. The
apophyses are no doubt useful in preserving the delicate vital organs
from shocks, or from derangement by the muscular movements and
the changes of position of the body.
Fig. 57.—Transverse section of skeleton of metathorax of Goliathus

druryi, seen from behind: a, metanotum; b, metasternum; c,
phragma; d, entothorax (apophysis or furca); e, apodeme; f,
tendon of articulation. (After Kolbe.)
The appendages of the thorax are (a) inferior, the legs; (b) superior,
the wings. The legs are always six in number, and are usually
present even in larvae, though there exist many apodal larvae,
especially in Diptera. The three pairs of legs form one of the most
constant of the characters of Insects. They are jointed appendages
and consist of foot, otherwise tarsus; tibia, femur, trochanter, and
coxa; another piece, called trochantin more or less distinctly
separated from the coxa, exists in many Insects. The legs are
prolongations of the body sac, and are in closer relation with the
epimera and with the episterna than with other parts of the crust,
though they have a close relation with the sternum. If we look at the
body and leg of a neuropterous Insect (Fig. 58) we see that the basal
part of the leg—the coxa—is apparently a continuation of one of the
two pleural pieces or of both; in the latter case one of the prolonged
pieces forms the coxa proper, and the tip of the other forms a
supporting piece, which may possibly be the homologue of the
trochantin of some Insects. In some Orthoptera, especially in
Blattidae, and in Termitidae, there is a transverse chitinised fold
interposed between the sternum and the coxa, and this has the
appearance of being the same piece as the trochantin of the anterior
legs of Coleoptera.
Fig. 58.—Hind leg of Panorpa: a, episternum; a′, epimeron; b, coxa; b′,

coxal fold of epimeron; c, trochanter; d, femur; e, tibia; f, tarsus.
Beyond the coxa comes the trochanter; this in many Hymenoptera is

a double piece, though in other Insects it is single; usually it is the
most insignificant part of the leg. The femur is, on the whole, the
least variable part of the leg; the tibia, which follows it, being
frequently highly modified for industrial or other purposes. The joint
between the femur and the tibia is usually bent, and is therefore the
most conspicuous one in the leg; it is called the knee. The other
joints have not corresponding names, though that between the tibia
and the tarsus is of great importance. The spines at the tip of the
tibia, projecting beyond it, are called spurs, or calcares. The tarsus
or foot is extremely variable; it is very rarely absent, but may consist
of only one piece—joint, as it is frequently called[23]—or of any larger
number up to five, which may be considered the characteristic
number in the higher Insect forms. The terminal joint of the tarsus
bears normally a pair of claws; between the claws there is frequently
a lobe or process, according to circumstances very varied in different
Insects, called empodium, arolium, palmula, plantula,
pseudonychium, or pulvillus. This latter name should only be used in
those cases in which the sole of the foot is covered with a dense
pubescence. The form of the individual tarsal joints and the armature
or vestiture of the lower surface are highly variable. The most
remarkable tarsus is that found on the front foot of the male
Dytiscus.
It has been suggested that the claws and the terminal appendage of
the tarsus ought to be counted as forming a distinct joint; hence
some authors state that the higher Insects have six joints to the feet.
These parts, however, are never counted as separate joints by
systematic entomologists, and it has recently been stated that they
are not such originally.
The parts of the foot at the extremity of the last tarsal joint proper are
of great importance to the creature, and vary greatly in different
Insects. The most constant part of this apparatus is a pair of claws,
or a single claw. Between the two claws there may exist the
additional apparatus referred to above. This in some Insects—
notably in the Diptera—reaches a very complex development. We
figure these structures in Pelopaeus spinolae, a fossorial
Hymenopteron, remarking that our figures exhibit the apparatus in a
state of retraction (Fig. 59). According to the nomenclature of Dahl
and Ockler[24] the plate (b) on the dorsal aspect is the pressure plate
(Druck-Platte), and acts as an agent of pressure on the sole of the
pad (C, e); c and d on the underside are considered to be extension-
agents; c, extension-plate; d, extension-sole (Streck-Platte, Streck-
Sohle). These agents are assisted in acting on the pad by means of
an elastic bow placed in the interior of the latter. The pad (e) is a
very remarkable structure, capable of much extension and retraction;
when extended it is seen that the pressure plate is bent twice at a
right angle so as to form a step, the distal part of which runs along
the upper face of the basal part of the pad; the apical portion of this
latter consists of two large lobes, which in repose, as shown in our
Figure (f), fall back on the pad, something in the fashion of the
retracted claws of the cat, and conceal the pressure-plate.
The mode in which Insects are able to walk on smooth perpendicular

surfaces has been much discussed, and it appears highly probable
that the method by which this is accomplished is the exudation of
moisture from the foot; there is still, however, much to be ascertained
before the process can be satisfactorily comprehended. The theory
to the effect that the method is the pressure of the atmosphere
acting on the foot when the sole is in perfect apposition with the
object walked on, or when a slight vacuum is created between the
two, has apparently less to support it.
Fig. 59.—Foot of Pelopaeus, a fossorial wasp: A, tarsus entire; B,

terminal joint, upper side; C, under side. a, claw; b, base of
pressure-plate; c, extension-plate; d, extension-sole; e, pad; f,
lobe of pad retracted.
The legs of the young Insect are usually more simple than those of
the adult, and in caterpillars they are short appendages, and only
imperfectly jointed. If a young larva, with feet, of a beetle, such as
Crioceris asparagi be examined, it may be seen that the leg is
formed by protuberance of the integument, which becomes divided
into parts by simple creases; an observation suggesting that the
more highly developed jointed leg is formed in a similar manner. This
appears to be really the case, for the actual continuity of the limb at
the chief joint—the knee—can be demonstrated in many Insects by
splitting the outer integument longitudinally and then pulling the
pieces a little apart; while in other cases even this is not necessary,
the knee along its inner face being membranous to a considerable
extent, and the membrane continuous from femur to tibia.
Turning to the wings, we remark that there may be one or two pairs
of these appendages. When there is but one pair it is nearly always
mesothoracic, when there are two pairs one is invariably
mesothoracic, the other metathoracic. The situation of the wing is
always at the edge of the notum, but the attachment varies in other
respects. It may be limited to a small spot, and this is usually the
case with the anterior wing; or the attachment may extend for a
considerable distance along the edge of the notum, a condition
which frequently occurs, especially in the case of the posterior
wings. The actual connexion of the wings with the thorax takes place
by means of strong horny lines in them which come into very close
relation with the little pieces in the thorax which we have already
described, and which were styled by Audouin articulatory epidemes.
There is extreme variety in the size, form, texture, and clothing of the
wings, but there is so much resemblance in general characters
amongst the members of each one of the Orders, that it is usually
possible for an expert, seeing only a wing, to say with certainty what
Order of Insects its possessor belonged to. We shall allude to these
characters in treating of the Orders of Insects.
Each wing consists of two layers, an upper and a lower, and

between them there may be tracheae and other structures,
especially obvious when the wings are newly developed. It has been
shown by Hagen that the two layers can be separated when the
wings are recently formed, and it is then seen that each layer is
traversed by lines of harder matter, the nervures. These ribs are
frequently called wing-veins, or nerves, but as they have no relation
to the anatomical structures bearing those names, it is better to
make use of the term nervures. The strength, number, form and
inter-relations of these nervures vary exceedingly; they are thus
most important aids in the classification of Insects. Hence various
efforts have been made to establish a system of nomenclature that
shall be uniform throughout the different Orders, but at present
success has not attended these efforts, and it is probable that no
real homology exists between the nervures of the different Orders of
Insects. We shall not therefore discuss the question here. We may,
however, mention that German savants have recently distinguished
two forms of nervures which they consider essentially distinct, viz.
convex and concave. These, to some extent, alternate with one
another, but a fork given off by a convex one is not considered to be
a concave one. The terms convex and concave are not happily
chosen; they do not refer to the shape of the nervures, but appear to
have been suggested by the fact that the surface of the wing being
somewhat undulating the convex veins more usually run along the
ridges, the concave veins along the depressions. The convex are the
more important of the two, being the stronger, and more closely
connected with the articulation of the wing.
The wings, broadly speaking, may be said to be three-margined: the

margin that is anterior when the wings are extended is called the
costa, and the edge that is then most distant from the body is the
outer margin, while the limit that lies along the body when the wings
are closed is the inner margin.
The only great Order of Insects provided with a single pair of wings
is the Diptera, and in these the metathorax possesses, instead of
wings, a pair of little capitate bodies called halteres or poisers. In the
abnormal Strepsiptera, where a large pair of wings is placed on the
metathorax, there are on the mesothorax some small appendages
that are considered to represent the anterior wings. In the great
Order Coleoptera, or beetles, the anterior wings are replaced by a
pair of horny sheaths that close together over the back of the Insect,
concealing the hind-wings, so that the beetle looks like a wingless
Insect: in other four-winged Insects it is usually the front wings that
are most useful in flight, but the elytra, as these parts are called in
Coleoptera, take no active part in flight, and it has been recently
suggested by Hoffbauer[25] that they are not the homologues of the
front wings, but of the tegulae (see Fig. 56), of other Insects. In the
Orthoptera the front wings also differ in consistence from the other
pair over which they lie in repose, and are called tegmina. There are
many Insects in which the wings exist in a more or less rudimentary
or vestigial condition, though they are never used for purposes of
flight.
The abdomen, or hind body, is the least modified part of the body,
though some of the numerous rings of which it is composed may be
extremely altered from the usual simple form. Such change takes
place at its two extremities, but usually to a much greater extent at
the distal extremity than at the base. This latter part is attached to
the thorax, and it is a curious fact that in many Insects the base of
the abdomen is so closely connected with the thorax that it has all
the appearance of being a portion of this latter division of the body;
indeed it is sometimes difficult to trace the real division between the
two parts. In such cases a further differentiation may occur, and the
part of the abdomen that on its anterior aspect is intimately attached
to the thorax may on its posterior aspect be very slightly connected
with the rest of the abdomen. Under such circumstances it is difficult
at first sight to recognise the real state of the case. When a segment
is thus transferred from the abdomen to the metathorax, the part is
called a median segment. The most remarkable median segment
exists in those Hymenoptera which have a stalked abdomen, but a
similar though less perfect condition exists in many Insects. When
such a union occurs, it is usually most complete on the dorsal
surface, and the first ventral plate may almost totally disappear: such
an alteration may involve a certain amount of change in the sclerites
of the next segment, so that the morphological determination of the
parts at the back of the thorax and front of the abdomen is by no
means a simple matter. A highly modified hind-body exists in the
higher ants, Myrmicidae. In Fig. 60 we contrast the simple abdomen
of Japyx with the highly modified state of the same part in an ant.
Fig. 60.—Simple abdomen of Japyx (A) contrasted with the highly
modified one of an ant, Cryptocerus (B). The segments are
numbered from before backwards.
Unlike the head and thorax, the abdomen is so loosely knitted

together that it can undergo much expansion and contraction. This is
facilitated by an imbricated arrangement of the plates, and by their
being connected by means of membranes admitting of much
movement (Fig. 47, m, p. 88). In order to understand the structure of
the abdomen it should be studied in its most distended state; it is
then seen that there is a dorsal and a ventral hard plate to each ring,
and there is also usually a stigma; there may be foldings or plications
near the line of junction of the dorsal and ventral plates, but these
margins are not really distinct pieces. The pleura, in fact, remain
membranous in the abdominal region, contrasting strongly with the
condition of these parts in the thorax. The proportions of the plates
vary greatly; sometimes the ventral are very large in proportion to the
dorsal, as is usually the case in Coleoptera, while in the Orthoptera
the reverse condition prevails.
Cerci or other appendages frequently exist at the extremity of the

abdomen (Fig. 47, n, p. 88); the former are sometimes like antennae,
while in other cases they may be short compressed processes
consisting of very few joints. The females of many Insects possess
saws or piercing instruments concealed within the apical part of the
abdomen; in other cases an elongate exserted organ, called
ovipositor, used for placing the eggs in suitable positions, is present.
Such organs consist, it is thought, either of modified appendages,
called gonapophyses, or of dorsal, ventral, or pleural plates. The
males frequently bear within the extremity of the body a more or less
complicated apparatus called the genital armour. The term
gonapophysis is at present a vague one, including stings, some
ovipositors, portions of male copulatory apparatus, or other
structures, of which the origin is more or less obscure.
The caterpillar, or larva, of the Lepidoptera and some other Insects,

bears a greater number of legs than the three pairs we have
mentioned as being the normal number in Insects, but the posterior
feet are in this case very different from the anterior, and are called
false legs or prolegs. These prolegs, which are placed on the hind
body, bear a series of hooks in Lepidopterous larvae, but the
analogous structures of Sawfly larvae are destitute of such hooks.
Placed along the sides of the body, usually quite visible in the larva,
but more or less concealed in the perfect Insect, are little apertures
for the admittance of air to the respiratory system. They are called
spiracles or stigmata. There is extreme variety in their structure and
size; the largest and most remarkable are found on the prothorax of
Coleoptera, especially in the groups Copridae and Cerambycidae.
The exact position of the stigmata varies greatly, as does also their
number. In the Order Aptera there may be none, while the maximum
number of eleven pairs is said by Grassi[26] to be attained in Japyx
solifugus: in no other Insect have more than ten pairs been recorded,
and this number is comparatively rare. Both position and number
frequently differ in the early and later stages of the same Insect. The
structure of the stigmata is quite as inconstant as the other points we
have mentioned are.
Fig. 61.—Membranous space between pro- and meso-thoraces of a

beetle Euchroma, showing stigma (st); a, hind margin of
pronotum; b, front leg; c, front margin of mesonotum; d, base of
elytra; e, mesosternum.
The admission of air to the tracheal system and its confinement
there, as well as the exclusion of foreign bodies, have to be provided
for. The control of the air within the system is, according to
Landois[27] and Krancher,[28] usually accomplished by means of an
occluding apparatus placed on the tracheal trunk a little inside of the
stigma, and in such case this latter orifice serves chiefly as a means
for preventing the intrusion of foreign bodies. The occluding
apparatus consists of muscular and mechanical parts, which differ
much in their details in different Insects. Lowne supposes that the air
is maintained in the tracheal system in a compressed condition, and
if this be so, this apparatus must be of great importance in the Insect
economy. Miall and Denny[29] state that in the anterior stigmata of
the cockroach the valves act as the occluding agents, muscles being
attached directly to the inner face of the valves, and in some other
Insects the spiracular valves appear to act partially by muscular
agency, but there are many stigmata having valves destitute of
muscles. According to Lowne[30] there exist valves in the blowfly at
the entrance to the trachea proper, and he gives the following as the
arrangement of parts for the admission of air:—there is a spiracle
leading into a chamber, the atrium, which is limited inwardly by the
occluding apparatus; and beyond this there is a second chamber, the
vestibule, separated from the tracheae proper by a valvular
arrangement. He considers that the vestibule acts as a pump to
force the air into the tracheae.
Fig. 62.—Diagrammatic Insect to explain terms of position. A, apex; B,

base: 1, tibia; 2, last abdominal segment; 3, ideal centre.
Systematic Orientation.
Terms relating to position are unfortunately used by writers on
entomology in various, even in opposite senses. Great confusion
exists as to the application of such words as base, apex, transverse,
longitudinal. We can best explain the way in which the relative
positions and directions of parts should be described by reference to
Figure 62. The spot 3 represents an imaginary centre, situated
between the thorax and abdomen, to which all the parts of the body
are supposed to be related. The Insect should always be described
as if it were in the position shown in the Figure, and the terms used
should not vary as the position is changed. The creature is placed
with ventral surface beneath, and with the appendages extended,
like the Insect itself, in a horizontal plane. In the Figure the legs are,
for clearness, made to radiate, but in the proper position the anterior
pair should be approximate in front, and the middle and hind pairs
directed backwards under the body. The legs are not to be treated as
if they were hanging from the body, though that is the position they
frequently actually assume. The right and left sides, and the upper
and lower faces (these latter are frequently also spoken of as sides),
are still to retain the same nomenclature even when the position of
the specimen is reversed. The base of an organ is that margin that is
nearest to the ideal centre, the apex that which is most distant. Thus
in Fig. 62, where 1 indicates the front tibia, the apex (A) is broader
than the base (B); in the antennae the apex is the front part, while in
the cerci the apex is the posterior part; in the last abdominal
segment (2) the base (B) is in front of the apex (A). The terms
longitudinal and transverse should always be used with reference to
the two chief axes of the body-surface; longitudinal referring to the
axis extending from before backwards, and transverse to that going
across, i.e. from side to side.
CHAPTER IV
ARRANGEMENT OF INTERNAL ORGANS–MUSCLES–NERVOUS SYSTEM–

GANGLIONIC CHAIN–BRAIN–SENSE-ORGANS–ALIMENTARY CANAL–
MALPIGHIAN TUBES–RESPIRATION–TRACHEAL SYSTEM–FUNCTION OF
RESPIRATION–BLOOD OR BLOOD-CHYLE–DORSAL VESSEL OR HEART–
FAT-BODY–OVARIES–TESTES–PARTHENOGENESIS–GLANDS.
The internal anatomy of Insects may be conveniently dealt with

under the following heads:—(1) Muscular system; (2) nervous
system; (3) alimentary system (under which may be included
secretion and excretion, about which in Insects very little is known);
(4) respiratory organs; (5) circulatory system; (6) fat-body; (7)
reproductive system.
Fig. 63.—Diagram of arrangement of some of the internal organs of an

Insect: a, mouth; b, mandible; c, pharynx; d, oesophagus; e,
salivary glands (usually extending further backwards); f, eye; g,
supra-oesophageal ganglion; h, sub-oesophageal ganglion; i,
tentorium; j, aorta; k1, k2, k3, entothorax; l1-l8, ventral nervous
chain; m, crop; n, proventriculus; o, stomach; p, Malpighian tubes;
q, small intestine; r, large intestine; s, heart; t, pericardial septum;
u, ovary composed of four egg-tubes; v, oviduct; w, spermatheca
(or an accessory gland); x, retractile ovipositor; y, cercus; z,
labrum.
Many of the anatomical structures have positions in the body that are
fairly constant throughout the class. Parts of the respiratory and
muscular systems and the fat-body occur in most of the districts of
the body. The heart is placed just below the dorsal surface; the
alimentary canal extends along the middle from the head to the end
of the body. The chief parts of the nervous system are below the
alimentary canal, except that the brain is placed above the beginning
of the canal in the head. The reproductive system extends in the
abdomen obliquely from above downwards, commencing anteriorly

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Smith & Tanagho’s General Urology

19th Edition Edition Jack W. Mcaninch

Smith & Tanagho’s General Urology 19th Edition Edition

Campbell Walsh Wein Urology 12th Edition Alan W Partin

Vaughan & Asbury's General Ophthalmology 19th Edition

Vaughan & Asbury’s General Ophthalmology 19th Edition

General, Organic, & Biological Chemistry 5th Edition

Campbell Walsh Wein Handbook of Urology 1st Edition

Principles of General, Organic, & Biological Chemistry

General, Organic, and Biological Chemistry 5th Edition

Smith & Tanagho’s

Jack W. McAninch, MD, FACS, FRCS(E)(Hon)

Tom F. Lue, MD, FACS, ScD (Hon)

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20 Renal Parenchymal Neoplasms 329 31 Disorders of the Adrenal Glands 509

21 Cancer of the Prostate Gland 351 32 Disorders of the Kidneys 521

22 Genital Tumors 377 33 Diagnosis of Medical Renal Diseases 539

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42 Disorders of the Female Urethra 659 46 Genital Gender-Affirming Surgery:

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Peter R. Carroll, MD, MPH Arpita Desai, MD

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Michael DiSandro, MD Kirsten L. Greene, MD, MS

Daniela Franz, MD Hedvig Hricak, MD, PhD

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John N. Krieger, MD Anobel Y. Odisho, MD, MPH

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Katsuto Shinohara, MD David Tat, DO

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Jack W. McAninch, MD, FACS, FRCS(E) (Hon)

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CALICES, RENAL PELVIS, AND URETER

1. Calices—The tips of the minor calices (8–12 in number)

▶▶Histology (Figure 1–4) ▶▶Gross Appearance

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B. Relations ▶▶Histology (Figure 1–10)

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▲▲Figure 1–11. Section of the prostate gland shows the

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▶▶Blood Supply EPIDIDYMIS

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The testis is covered anteriorly and laterally by the visceral SCROTUM

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▶▶Blood Supply ▶▶Blood Supply

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Emil A. Tanagho, MD; Hiep T. Nguyen, MD;

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