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Embriología del

Sistema
Nervioso
MSc. María Gabriela Trejo Sarmiento
Universidad Francisco Marroquín
Guatemala, Enero 2020

0
Objetivos
• Delinear el desarrollo embrionario inicial que lleva la formación del
sistema nervioso

• Describir la embriología del sistema nervioso en general y en


específico.

• Relacionar las anomalías embrionarias que llevan a patología en el


recién nacido y adulto

1
ÍNDICE
1. Introducción 4. Desarrollo de la médula espinal
a. Desarrollo humano a. Generalidades
b. Embriología b. Desarrollo de los ganglios espinales
c. Aspectos históricos c. Desarrollo de los meninges
espinales
2. Resumen
a. Primera semana d. Cambios posicionales
b. Segunda semana e. Mielinización de las fibras nerviosas
c. Tercera semana f. Defectos en el desarrollo de la
médula espinal
3. Desarrollo del sistema nervioso
a. Generalidades 5. Desarrollo del cerebro
b. Embriología del sistema nervioso a. Generalidades
c. Defectos en el desarrollo del SN b. Defectos en el desarrollo del
cerebro
2
morphologic characteristics. Stage 1 begins at fertiliza- rapidly during infancy; total length increases by approxi-
tion and embryonic development ends at stage 23, which mately one half and weight is usually tripled. By 1 year
occurs on day 56 (see Fig. 1-1). A trimester is a period of age, most infants have six to eight teeth.
of 3 months, one third of the 9-month period of gesta-
tion. The most critical stages of development occur during
the first trimester (13 weeks), when embryonic and early Childhood
fetal development is occurring. This is the period between infancy and puberty. The
primary (deciduous) teeth continue to appear and are
later replaced by the secondary (permanent) teeth. During
Postnatal Period early childhood, there is active ossification (formation of
This is the period occurring after birth. Explanations of bone), but as the child becomes older, the rate of body
frequently used developmental terms and periods follow. growth slows down. Just before puberty, however, growth
accelerates—the prepubertal growth spurt.
Infancy

Introducción: Desarrollo Humano


This is the period of extrauterine life, roughly the first Puberty
year after birth. An infant age 1 month or younger This is the period when humans become functionally
is called a neonate. Transition from intrauterine to capable of procreation (reproduction). Reproduction is

TIMETABLE OF HUMAN PRENATAL DEVELOPMENT


1 TO 10 WEEKS

El desarrollo humano es un proceso


Primary follicles Oocyte

EARLY DEVELOPMENT OF OVARIAN FOLLICLE

continuo que comienza cuando un


oocito es fertilizado por un
MENSTRUAL PHASE PROLIFERATIVE PHASE
Day 1 of last normal
menstrual cycle

espermatozoide y forman un cigoto. Antrum

COMPLETION OF DEVELOPMENT OF FOLLICLE


Mature
follicle
Oocyte
Ovulation

Oocyte Oocyte Ovary


CONTINUATION OF PROLIFERATIVE PHASE OF MENSTRUAL CYCLE

AGE
(weeks) 1 Stage 1 2 Stage 2 begins 3 4 Stage 3 begins 5 6 Stage 4 7 Stage 5 begins
Trophoblast
Zona pellucida Implantation begins

Fertilization Zygote divides Morula Early blastocyst Late blastocyst


Embryoblast
SECRETORY PHASE OF MENSTRUAL CYCLE

8 9 10 Cytotrophoblast 11 Maternal blood 12 Extraembryonic 13 Stage 6 begins 14


Lacunar Connecting stalk
Lacunae appear in Amnion Eroded mesoderm
syncytiotrophoblast gland network Primary villi Amnion
Amniotic cavity

Primary Embryonic disc


Bilaminar embryonic Primary umbilical umbilical Coelom
disc vesicle Closing plug vesicle Embryonic disc Prechordal plate

F I G U R E 1 – 1 Early stages of development. Development of an ovarian follicle containing an oocyte, ovulation, and the phases
of the menstrual cycle are illustrated. Human development begins at fertilization, approximately 14 days after the onset of the last
normal menstrual period. Cleavage of the zygote in the uterine tube, implantation of the blastocyst in the endometrium3(lining) of the
uterus, and early development of the embryo are also shown. The alternative term for the umbilical vesicle is the yolk sac; this is an
inappropriate term because the human vesicle does not contain yolk.
Introducción: Desarrollo Humano

División celular
Migración celular
Apoptosis Organismo
Diferenciación celular multicelular
Crecimiento celular
Reordenamiento celular

Célula totipotente
altamente especializada 4
Introducción: Desarrollo Humano
• La mayoría de cambios ocurren durante los períodos embrionario y fetal;
sin embargo se producen cambios importantes durante los últimos CHAPTER 1 |
|
períodos de desarrollo
CHAPTER 1 INTRODUCTION TO HUMAN DEVELOPMENT 9
Superior
Superior

• El desarrollo NO se detiene al nacer Cranial


Cranial

Dorsal Anterior Posterior


Anterior Posterior

Ventral Ventr

Caudal
Inferior
A B Inferior 5
A B
Sagittal plane
Sagittal plane

Lateral

Lateral
Amniotic sac
Wall of
uterus
Head large but chin Eyelid
Uterine Urogenital
poorly formed.
cavity membrane
Grooves between
digital rays External ear
Anal
indicate fingers. membrane Wrist,
Eyelids fingers
CRL: 13.0 mm forming Smooth or fused CRL: 18 mm
chorion

F I G U ed into an derived from rm in penet


transf
0 Stage 20 begins 51 52 Stage 21 begins 53 54 Stage 22 begins 55 56 Stage 23

some
ning o
Introducción: Desarrollo Humano Genital

orm
Ear

. The
Eye tubercle

RE 2
f fertil
Upper limbs Ear
longer and bent Eye

acroso
External genitalia Urethral

iz
at elbows. Wrist

Nucle d
groove

– 4 Illu ated sper e Golgi reg g the coro


have begun

a
cover some
by acr

t
Nose

FIGU

io
of the
to differentiate.

region

Head

me,
huma

n
Fingers distinct

Middle
of tail

us
Fingers

elong
e
Knee

t
The ep
• El desarrollo de un humano desde la fertilización de un oocitoElbow
hasta el
The e transports t re free-swi (Fig. 2-5A and tail. T m

o
o
which ure sperms d and a ta een the hea of the spe
but webbed. Anus

stratio
consis

a
the sp f the sperm

A
n seco

nucleu
s, the ary oocyte

Acros
head

RE 2

Neck

ssist t
16 THE DEVELOPING HUMAN

Mat

p
nacimiento esta dividida en dos períodos:

iece
Toes

pididy
or

ididym

ns of
Large forehead Toes

o
ting o the junctio most of th

he sp
nd
CRL: 30 mm

s, is p
erm is
o

middle
Golgi region

me
Acrosome Residual cytoplasm

– 5 M covered by piece, and e e zona pellu

Mitoch
Nucleus
7 58 59 60 61 62 Genitalia 63

sperm
Placenta
mis is Genitalia

m. No
e
is is an

t
a
f a he

h
r
Phallus

t
piece

ale an
Phallus

ly

Princip

ondria
Ear

iogen
Centrioles
Eye

te the
• Embrionario: ocurren los principales avances, de la tercera a la octava semana de
Urogenital
he spe

Mitochondrion
contin
a

Urogenital
a

, princ
elonga us with th ra (see Fig. cells

(×200
Beginning

d fe
fold

l shea
al piec
fold

e
forms

Nucleus Acrosome
of
embarazo)

sis, th
Wrist

loss o
r
male
rms to

a
Labioscrotal

ion (fi
), surr
uo

fetal

ip
n betw

th
in
Labioscrotal
ted co

e of ta
a
period Knee fold

e last
l
the ca

f cyto
fold

End p
game

rst dra a radiata a


o
mmin

u
the ur
il

Mitochondrial sheath
Perineum

il
nded
Perineum
iled du

F I G U R E 2 – 4 Illustrations of spermiogenesis, the last phase of spermatogenesis. During this process, the rounded spermatid is

p
p
• Fetal: Se produce la diferenciación y el crecimiento de tejidos y órganos, aumentan
p-li

iece o
tes (se osome, an
transformed into an elongated sperm. Note the loss of cytoplasm (see Fig. 2-5C), development of the tail, and formation of the acro-
Elbow

h
la
wing)
Toes

ase o
some. The acrosome, derived from the Golgi region (first drawing) of the spermatid, contains enzymes that are released at the begin-
g, acti

sm (se
CRL: 45 mm
ke acr
eth

n
ning of fertilization to assist the sperm in penetrating the corona radiata and zona pellucida surrounding the secondary oocyte.
b

CRL: 50 mm
e duct

las tasas de crecimiento corporal.


y

f tail
e bulk

ct (see deferens,

x cells
t

f sper
4 65 66 67 68 69 70

of the d zona pell


nd pie
h

Clitoris

e Fig.
vely m

Follicular cells of
)
d

Acrosome
.

us

corona radiata
Glans of penis
). A, T

Principal piece of tail

matog developme zymes that secondary


Fig. 2-
T

Labium

s
ce.

Face has

p
h

2
Head
Genitalia have First polar body

ermat
otile

minus

-
e neck e

5
more developed
B, A s

Nucleus

C
or Urethral
he ma

enesis
2-12).

covered

),
profile.
organ
1

by acrosome
cida a

Urogenital characteristics groove


2

id, co
Cytoplasm
)

Neck
groove
perm
.

but still not


o

in par taining enz


h
r

. Durin
Note growth
elle c

Nucleus
f

fully formed.
nd c

n
ucida
of chin
Middle piece
Scrotum

tains e unding the


Labium
and co s covered g severa

of tail
drawn
head le containi sed, the
organ When rele f the co

ts of a

compared
2-5A) cular cells pellucid

orona

on

Follicu
of foll of the zona -13A a

g this
c
majus
t

nt of t
2-5A

o
ration d C and 2

Ears still lower 6

surro
Zona
to day 44.

r
pellucida

n
n
than normal.
i

t
n
e

CRL: 61 mm

a
C

lar ce
o
.

huma
radiat

proce
End piece of tail
l

t
i

he tail
ains th
an

a
appro

diata
A B C
FIGURE 1–1, cont’d

lls of
n spe
a.

F I G U R E 2 – 5 Male and female gametes (sex cells). A, The main parts of a human sperm (×1250). The head, composed mostly

ss, the rmation o


ymes.

, and
of the nucleus, is partly covered by the cap-like acrosome, an organelle containing enzymes. The tail of the sperm consists of three
ximat

regions, the middle piece, principal piece, and end piece. B, A sperm drawn to approximately the same scale as the oocyte. C, A
e

First p

are re
human secondary oocyte (×200), surrounded by the zona pellucida and corona radiata.
rm (×1 il of
n
b
u

round
fo
n

ely th
a

The ta m
y the
o

cleus. c

olar b

leased
The epididymis is an elongated coiled duct (see Fig. 2-12). and contains the nucleus. The anterior two thirds of the
250).

The epididymis is continuous with the ductus deferens, head is covered by the acrosome, a cap-like saccular
e sa

ed sp
ody
which transports the sperms to the urethra (see Fig. 2-12). organelle containing several enzymes (see Figs. 2-4 and
Mature sperms are free-swimming, actively motile cells 2-5A). When released, the enzymes facilitate dispersion
T
a

a
T
h

o
Introducción: Embriología
Clínicamente orientada se refiere al estudio de embriones; estudia los
cambios estructurales de un ser humano desde la fertilización hasta el
nacimiento.

• Conocimientos sobre los comienzos de la vida y cambios que ocurren


durante el desarrollo prenatal.
• Permite la comprensión de las causas de las variaciones en estructura
humana
• Ilumina la anatomía orientada clínicamente
• Apoya la investigación y aplicación de células madre para tratamiento de
enfermedades crónicas.
7
Aspectos históricos
Egipcios (3000 a.C.):
• Métodos para incubar huevos de aves
• Atón à creador: germen en la mujer y semilla en el hombre
• El alma entraba al bebé al nacer a través de la placenta.
Indios (1416 a.C.):
• Tratado de embriología en Sanscrito (Garbha Upanishad)
• Período de concepción después de la conjugación de la sangre y el semen
• Describen diferentes estadios embrionarios y fetales
Griegos
• Hipócrates (460-377 a.C.): Incubación de huevos de gallina
• Aristóteles (284-322 a.C): Semen como la “semilla primordial” que daba lugar al embrión a través del semen y
la sangre menstrual.
• Galeno (130-201 d.C): descripción de la formación del feto, alantoides, amnios y placenta.

8
Aspectos históricos CHAPTER 1

Take twenty or more eggs and let them be incubated by


| INTRODUCTION TO HUMAN DEVELOPMENT 5

two or more hens. Then each day from the second to


that of hatching, remove an egg, break it, and examine
it. You will find exactly as I say, for the nature of the
bird can be likened to that of man.

Judíos Aristotle of Stagira (circa 384–322 BC), a Greek phi-


losopher and scientist, wrote a treatise on embryology in
• Talmud: 6 fases de formación del embrios which he described development of the chick and other A B
embryos. Aristotle promoted the idea that the embryo
developed from a formless mass, which he described as a
“less fully concocted seed with a nutritive soul and all
bodily parts.” This embryo, he thought, arose from men-
Edad Media strual blood after activation by male semen.
Claudius Galen (circa 130–201 AD), a Greek physi-
• Corán (Siglo VII d.C.): mezcla de secreciones del hombre y la mujer dan lugar
cian and medical scientist in Rome, wrote a book,
On the Formation of the Foetus, in which he described
a un nuevo organismo the development and nutrition of fetuses and the
structures that we now call the allantois, amnion, and
C D E

• Constantino el Africano (1020-1087 d.C): De Humana Natura , donde


placenta.
The Talmud contains references to the formation of
describió la composición y desarrollo secuencial del embrión en relación con
the embryo. The Jewish physician Samuel-el-Yehudi, who
lived during the second century AD, described six stages
los planetas y cada mes del embarazo. in the formation of the embryo from a “formless, rolled-up
thing” to a “child whose months have been completed.”
• Esquemas de fetos: Niños plenamente desarrollados en el interior del útero.
Talmud scholars believed that the bones and tendons, the
nails, the marrow in the head, and the white of the eyes,
were derived from the father, “who sows the white,” but
the skin, flesh, blood, and hair were derived from the F G
mother, “who sows the red.” These views were according
to the teachings of both Aristotle and Galen. F I G U R E 1 – 2 A-G, Illustrations from Jacob Rueff’s De Con-
ceptu et Generatione Hominis (1554) showing the fetus develop-
ing from a coagulum of blood and semen in the uterus. This
Embryology in the Middle Ages theory was based on the teachings of Aristotle, and it survived
The growth of science was slow during the medieval until the late 18th century. (From Needham J: A history of embry-
period, but a few high points of embryologic investiga- ology, ed 2, Cambridge, United Kingdom, 1934, Cambridge Uni-
tion undertaken during this time are known to us. It is versity Press; with permission of Cambridge University Press,
cited in the Quran (seventh century AD), the Holy Book England.)
of Islam, that human beings are produced from a mixture 9
of secretions from the male and female. Several references (Fig. 1-3). He introduced the quantitative approach to
are made to the creation of a human being from a nutfa embryology by making measurements of prenatal growth.
(small drop). It also states that the resulting organism It has been stated that the embryologic revolution
settles in the womb like a seed, 6 days after its beginning. began with the publication of William Harvey’s book De
Reference is made to the leech-like appearance of the Generatione Animalium in 1651. Harvey believed that
early embryo. Later the embryo is said to resemble a the male seed or sperm, after entering the womb or
“chewed substance.” uterus, became metamorphosed into an egg-like sub-
Constantinus Africanus of Salerno (circa 1020–1087 stance from which the embryo developed. Harvey (1578–
AD) wrote a concise treatise entitled De Humana Natura. 1657) was greatly influenced by one of his professors at
Africanus described the composition and sequential the University of Padua, Fabricius of Acquapendente, an
development of the embryo in relation to the planets and Italian anatomist and embryologist who was the first to
6 THE DEVELOPING HUMAN

Aspectos históricos
Renacimiento
• Leonardo da Vincci (1452-1519): Dibujos de disecciones de úteros
gestantes, introducción del método cuantitativo (mediciones de
crecimiento prenatal). F I G U R E 1 – 3 Reproduction of Leonardo da Vinci’s drawing
made in the 15th century showing a fetus in a uterus that has

• William Harvey (1578-1657): Revolución de la embriología con su libro been incised and opened.

De Generatione Animalium. Espermatozoide = semilla que en el útero se


convierte en ”huevo” Graaf observed small chambers in the rabbit’s uterus and
concluded that they could not have been secreted by the
• Girodamo Fabricius (1537-1619): Primer atlas y tratado De formato Foetu uterus. He stated that they must have come from organs
that he called ovaries. Undoubtedly, the small chambers
that de Graaf described were blastocysts (see Fig. 1-1).
• MICROSCOPIO He also described follicles which were called graafian
follicles; they are now called vesicular ovarian follicles.
• Reginier de Graaf (1672): Examina úteros de conejo àBlastocisto en Marcello Malpighi, studying what he believed was
unfertilized hen’s eggs in 1675, observed early embryos.
folículos ováricos– folículos de Graaf As a result, he thought the egg contained a miniature
chick. A young medical student in Leiden, Johan

• Johan Ham y Anton Van Leeuwenhoek: Microscopio modificado, Ham van Arnheim, and his countryman Anton van
Leeuwenhoek, using an improved microscope in 1677,
observan espermatozoides humanos = ser humano miniatura. first observed human sperms. However, they misunder-
stood the sperm’s role in fertilization. They thought the
sperm contained a miniature preformed human being
that enlarged when it was deposited in the female genital
tract (Fig. 1-4).
Caspar Friedrich Wolff refuted both versions of the
preformation theory in 1759, after10 observing that parts
of the embryo develop from “globules” (small spherical
bodies). He examined unincubated eggs but could not see
the embryos described by Malpighi. He proposed the
layer concept, whereby division of what we call the zygote
produces layers of cells (now called the embryonic disc)
from which the embryo develops. His ideas formed the
basis of the theory of epigenesis, which states that “devel-
opment results from growth and differentiation of spe-
cialized cells.” These important discoveries first appeared
in Wolff’s doctoral dissertation Theoria Generationis. He
also observed embryonic masses of tissue that partly con-
Aspectos históricos
Renacimiento
• Caspar Wolff: concepto de desarrollo por capas – disco embrionario.
Observó masas de tejido embrionario que contribuyen en parte al
desarrollo de los sistemas urinario y genital!!
• Lazaro Spallanzani (1775): Inseminación artificial en perros à el oocito y
espermatozoide son necesarios para formar un nuevo individuo
• Heinrich Cristian Pander (1817): 3 capas germinales del embrión -
BLASTODERMO
• Etienne e Isidore Saint Hilaire: Estudian alteraciones del desarrollo,
indujeron malformaciones congénitas en animales, nace la teratología.
• Karl Ernst von Baer: Describe oocitos en los folículos ováricos. Padre de la
embriología moderna, observo divisiones. Describió las capas germinales
y sus derivados.

11
Aspectos históricos
• Matthias Schleiden y Theodor Schwann (1839): teoría celular – De
una única célula (cigoto) se divde para formar un embrión.
• Hans Spermann (1935):Fenómeno de la inducción primaria.
• Robert Edwards y Patrick Steptoe (1978): Fecundación In Vitro,
nacimiento de Louise Brown

12
Meiotic spindle Antrum oocyte
Zona pellucida
23,X in
Corona radiata mature follicle

23,X 23,X 23,Y 23,Y


Spermatids
SPERMIOGENESIS
Secondary oocyte First polar body
First polar body

A
First meiotic
division completed
Corona radiata
Pronucleus
Normal
sperms

Resumen (1era semana) 32

32
THE DEVELOPING HUMAN
Pronucleus
Sperm
DegeneratingSecond polar body
THE DEVELOPING HUMAN
tail of sperm
First and second
Second polar body

23,X 23,X 23,Y 23,Y Zona pellucida polar bodies


Second polar body
Second polar body
Second meiotic
B Fertilized
C oocyte division completed
Zona pellucida

1. Fertilización F I G U R E 2 – 1 Normal gametogenesis: conversion of germ cells into gametes (sex cells). The drawings compare spermatogenesis
and oogenesis. Oogonia are not shown in this figure, because they differentiate into primary oocytes before birth. The chromosome
Chromosomes
Blastomere Cleavage spindle

2. Formación del cigoto


A 2-cell stage Blastomere B 4-cell stage
complement of the germ cells is shown at each stage. The number designates the total number of chromosomes, including the sex Zygote
chromosome(s) shown after the comma. Notes: (1) Following the two meiotic divisions, the diploid number of chromosomes, 46, is
A 2-cell stage B 4-cell stage
reduced to the haploid number, 23. (2) Four sperms form from one primary spermatocyte, whereas only one mature oocyte results
Breakdown

3. Clivaje del cigoto of pronuclear


from maturation of a primary oocyte. (3) The cytoplasm is conserved during oogenesis to form one large cell, the mature oocyte (see
D
membranes
E
Fig. 2-5C). The polar bodies are small nonfunctional cells that eventually degenerate.
F I G U R E 2 – 1 4 Illustrations of fertilization, the procession of events beginning when the sperm contacts the secondary oocyte’s

4. Formación de una mórula Embrioblasto


plasma membrane and ending with the intermingling of maternal and paternal chromosomes at metaphase of the first mitotic division
of the zygote. A, Secondary oocyte surrounded by several sperms, two of which have penetrated the corona radiata. (Only 4 of the
23 chromosome pairs are shown.) B, The corona radiata is not shown. A sperm has entered the oocyte, and the second meiotic divi-

5. Formación del blastocisto Cavidad sion has occurred, causing a mature oocyte to form. The nucleus of the oocyte is now the female pronucleus. C, The sperm head has

fusing. E, The zygote has formed; it contains 46 chromosomes,


C the diploid
8-cell stage number.
Zona pellucida
enlarged to form the male pronucleus. This cell, now called an ootid, contains the male and female pronuclei. D, The pronuclei are
D Morula

Trofoblasto
Zona pellucida CHAPTER 2 | FIRST W

6. Formación de estructuras
C 8-cell stage D Morula Endometrial gland
signals (attractants), secreted by the oocyte and surround- chromosomes at metaphase of the first mitotic division
ing follicular cells, guide the capacitated sperms (sperm of the zygote, a unicellular
Embryoblast
embryo (see Fig. 2-14E).

extraembrionarias y la parte
chemotaxis) to the oocyte. Defects at any stage
(inner in the sequence of these events
cell mass)
Fertilization is a complex sequence of coordinated may cause the death of
Embryoblast
the zygote. The fertilization
molecular events that begins with contact between a process takes approximately
Degenerating
(inner cell mass) 24 hours. Transgenic and Endometrial

embrionaria de la placenta
capillary
sperm and an oocyte (see Fig. 2-13A and B) and gene knockout zona pellucida
studies in animals have shown that
ends with the intermingling of maternal and paternal carbohydrate-binding molecules and gamete-specific
Degenerating
zona pellucida
Blastocystic Endometrial
Embryonic epithelium
cavity

7. Adhesión al epitelio endometrial


pole FIG
Blastocystic to t
Embryoblast stag
cavity
Trophoblast
Blastocystic is at
emb
E Early blastocyst Trophoblast
cavity F Later blastocyst
sync
| of the blastocyst. A to D, Various stages of cleavage of the
CHAPTER 2
F I G U R E 2 – 1 6 Illustrations of a cleaving zygote and formation
FIRST WEEK OF HUMAN DEVELOPMENT Trophoblast
35

8. Diferenciación del trofoblasto en


and
E Early blastocyst A F Later blastocyst tive t
zygote. The period of the morula begins at the 12-cell to 16-cell stage and ends when the blastocyst forms. E and F, Sections of
Endometrial gland is us
F I G U RThe
blastocysts. E 2 –16
zona Illustrations
pellucida of a cleaving by
has disappeared zygote andblastocyst
the late formation stage
of the (5blastocyst.
days). TheA second
to D, Various stages of
polar bodies cleavage
shown in A of
arethesmall, Beca
zygote. The period of the morula begins at the 12-cell to 16-cell stage and ends when the blastocyst forms. E and F, Sections of

dos capas
nonfunctional cells. Cleavage of the zygote and formation of the morula occur as the dividing zygote passes along the uterine tube.
Endometrial
surfa
blastocysts. The zona pellucida has disappeared by the late blastocyst stage (5 days). The second polar bodies shown in A are small,tissue logic
Blastocyst formation occurs in the uterus. Although cleavage increases the number of blastomeres, note that each of the daughter connective
nonfunctional cells. Cleavage of the zygote and formation of the morula occur as the dividing zygote passes along the uterine tube. In th
cells is smaller than the parent cells. As a result, there is no increase in the size of the developing embryo until the zona pellucida when
Blastocyst formation occurs in the uterus. Although cleavage increases the number of blastomeres, note that each of the daughter
degenerates. The blastocyst then enlarges considerably (F).
cells is smaller than the parent cells. As a result, there is no increase in the size of the developing embryo until the zona pellucida
ation
Endometrial tion
capillary
degenerates. The blastocyst then enlarges considerably (F).

9. Implantación
inter
Glandular
secretion

Endometrial
Embryonic epithelium
Syncytiotrophoblast
pole F I G U R E 2 – 1 9 Attachment of the blastocyst
to the endometrial epithelium during the early Embryoblast
Embryoblast stages of implantation. A, At 6 days, the trophoblast
Blastocystic is attached to the endometrial epithelium at Cytotrophoblast
the
cavity embryonic pole of the blastocyst. B, At 7 days, the

A
Trophoblast
13
syncytiotrophoblast has penetrated the epithelium
and has started to invade the endometrial connec-
Hypoblast

B tissue. Note: In embryologic studies, the embryo


tive
Blastocystic cavity
is usually shown with its dorsal surface upward.
Because the embryo implants on its future dorsal
surface, it would appear upside down if the histo-
Endometrial
connective tissue enzymes
logic that (epithelium
convention erode the upward)
maternal tissues,
were enabling the
followed. ampulla, wh
blastocyst
In this book,tothe“burrow”
histologicinto the endometrium.
convention is followed Endome- oocyte is exp
trial cells
when also assist tois control
the endometrium the depth
the dominant of penetration
consider- ● Sperms are
of the
ation syncytiotrophoblast.
(e.g., At approximately
Fig. 2-6C), and the embryologic conven- 7 days, a and are store
layerisofused
tion the hypoblast
cells,when the embryo (primary
is the endoderm),
center of appears lation of sem
on the surface
interest, as in theof the embryoblast
adjacent illustrations.facing the blastocystic sperms in th
Glandular cavity (see Fig. 2-19B). Comparative embryologic data through the
secretion suggest that the hypoblast arises by delamination of blas- ● When an ooc
tomeres from the embryoblast. the second m
a mature ooc
Syncytiotrophoblast The nucleus
SUMMARY OF FIRST WEEK female pronu
Embryoblast ● After the sp
● Oocytes are produced by the ovaries (oogenesis) and sperm separa
Cytotrophoblast expelled from them during ovulation (Fig. 2-20). The the male pro
fimbriae of the uterine tubes sweep the oocyte into the tilization is
Hypoblast

B Blastocystic cavity
frequently used developmental terms and periods follow. growth slows down. Just before puberty, however, growth
accelerates—the prepubertal growth spurt.
Infancy
This is the period of extrauterine life, roughly the first Puberty
year after birth. An infant age 1 month or younger This is the period when humans become functionally
is called a neonate. Transition from intrauterine to capable of procreation (reproduction). Reproduction is

TIMETABLE OF HUMAN PRENATAL DEVELOPMENT

Resumen (2da semana)


1 TO 10 WEEKS

Primary follicles Oocyte

EARLY DEVELOPMENT OF OVARIAN FOLLICLE

1. Rápida proliferación y diferenciación del trofoblasto – cuando el blastocisto completa la implantación


en el endometrio uterino. MENSTRUAL PHASE PROLIFERATIVE PHASE
Day 1 of last normal
2. Reacción decidual
menstrual cycle
Antrum Mature Oocyte

3. Formación de vesículas umbilicales primarias y se desarrolla el mesodermo extraembrionario. follicle Ovulation

COMPLETION OF DEVELOPMENT OF FOLLICLE

4. Cavidad coriónica
Oocyte Oocyte Ovary
5. La vesícula umbilical primaria se vuelve más pequeña y desaparece gradualmente a medida que se
CONTINUATION OF PROLIFERATIVE PHASE OF MENSTRUAL CYCLE

desarrolla la vesícula umbilical secundaria.


AGE
1 Stage 1 2 Stage 2 begins 3 4 Stage 3 begins 5 6 Stage 4 7 Stage 5 begins
(weeks) Trophoblast
6. Aparece la cavidad amniotica
Zona pellucida
(entre citotrofoblasto y embrioblasto) Implantation begins
7.
1
El embrioblasto se diferencia en un disco bilaminar Epiblasto à Cavidad amniotica
Fertilization Zygote divides Morula Early blastocyst Late blastocyst Hipoblasto
8. Engrosamiento de hipoblasto à futura región craneal
SECRETORY PHASE OF MENSTRUAL CYCLE
Embryoblast

8 9 10 Cytotrophoblast 11 Maternal blood 12 Extraembryonic 13 Stage 6 begins 14


Lacunar Connecting stalk
Lacunae appear in Amnion Eroded mesoderm
syncytiotrophoblast gland network Primary villi Amnion
Amniotic cavity

Bilaminar embryonic Primary umbilical


Primary Embryonic disc 14
Closing plug umbilical Coelom
disc vesicle vesicle Embryonic disc Prechordal plate

F I G U R E 1 – 1 Early stages of development. Development of an ovarian follicle containing an oocyte, ovulation, and the phases
of the menstrual cycle are illustrated. Human development begins at fertilization, approximately 14 days after the onset of the last
normal menstrual period. Cleavage of the zygote in the uterine tube, implantation of the blastocyst in the endometrium (lining) of the
uterus, and early development of the embryo are also shown. The alternative term for the umbilical vesicle is the yolk sac; this is an
inappropriate term because the human vesicle does not contain yolk.
Resumen (3era semana)
1. El disco embrionario bilaminar se convierte en trilaminar durante la gastrulación. Empieza con la aparición
de la línea primitiva.
2. La línea primitiva resulta de la migración de células epiblásticas al plano medio del disco. Esta invaginación
da lugar a la migración ventral, lateral y craneal de células mesenquimales entre el epiblasto y el hipoblasto.
3. Cuando la línea primitiva comienza a producir células mesenquimales:
• Epiblasto = ectodermo embrionario
• Hipoblasto = endodermo embrionario
• Células mesenquimatosas = tercera capa germinal (mesodermo intraembrionario)
• Células del mesodermo migran a los bordes del disco embrionario donde se unen a la cubierta del mesodermo
extraembrionario.
4. Al final de esta semana el embrión es un ovoide plano. El mesodermo existe entre el ectodermo y el
endodermo del disco, excepto en la membrana orofaríngea, en el plano medio ocupado por la
C H A P T E R 1 | INTRODUCTION TO HUMAN DEVELOPMENT
notocorda y en la membrana cloacal. 3

15 16 Stage 7 begins 17 Trilaminar embryo 18 Stage 8 begins 19 20 Stage 9 begins 21 Neural


First missed groove
menstrual period Amnion Neural plate
Neural plate
Brain First pairs
Neural of somites
Primitive groove Neural groove
3 streak Primitive
Somite Somite
streak
Arrows indicate Primitive node
Primitive streak
migration of Migration of cells from
primitive streak Length: 1.5 mm Primitive streak
Thyroid gland begins
to develop
15
mesenchymal cells
22 Stage 10 begins 23 Rostral neuropore 24 Stage 11 begins 25 26 Stage 12 begins 27 Site of otic pit 28 Stage 13 begins
Otic (ear) pit
Heart bulge Fore-
brain
Heart Primordia Upper
begins of eye Rostral limb bud Pharyngeal
4 to beat and ear neuropore
closes arches
present

Caudal
2 pairs of 3 pairs of Indicates CRL = crown−
neuropore
Neural folds fusing pharyngeal arches pharyngeal arches actual size rump length CRL: 5.0 mm
29 30 31 32 Stage 14 begins 33 Stage 15 begins 34 Cerebral vesicles 35 Eye
Developing eye distinct
Eye Upper
Sistema nervioso
El sistema nervioso es un conjunto de Sistema nervioso central (SNC)
células especializadas en la conducción de • Encéfalo
señales eléctricas. • Médula espinal
• Protegido por: Cráneo y la columna
vertebral
Está formado por neuronas y células
Sistema nervioso periférico (SNP)
gliales. • Neuronas
• Nervios craneales
• Ganglios nerviosos
Sistema nervioso autónomo (SNA)
• Neuronas que inervan el músculo liso, el
músculo cardíaco, el epitelio glandular y las
combinaciones de estos tejidos.
16
Desarrollo del sistema nervioso
• Tercera semana – aparece la primera indicación del desarrollo del SN
380 THE DEVELOPING HUMAN
• Inicio del desarrollo de la placa neural y surco neural en la cara posterior del embrión
trilaminar. Oropharyngeal membrane
Neural plate

Neural plate
Neural Neural
Notochordal process groove fold

Level of
section B

Primitive knot Primitive streak


Notochordal plate In
B m
Cloacal membrane
A Neural groove
17

Neura
Rostral
Neural fold neuropore

Neural
groove
Levels of
380 THE DEVELOPING HUMAN

Oropharyngeal membrane
Neural plate

Neural plate Amnion


Neural Neural
Notochordal process groove fold

Level of
section B Wall of
umbilical
vesicle
380 THE DEVELOPING HUMAN
380 THE DEVELOPING HUMAN Primitive knot Primitive streak
Notochordal plate Intraembryonic
B

Desarrollo del sistema nervioso


mesoderm
Oropharyngeal membrane
380 THE DEVELOPING HUMAN Neural plate Cloacal membrane
Oropharyngeal membrane
Neural plate A Neural plate Amnion Neural groove
Neural Neural
NeuralOropharyngeal
plate membrane Amnion
Notochordal process groove
NeuralNeural
plate fold
Neural Neural crest
groove Rostral
Notochordal process Neural foldfold neuropore
Neural plate Amnion
Neural Neural
Notochordal process groove Neural
fold
Level of groove

• Tubo neural à Sistema nervioso central


Levels of
section
Level of B Wall of
sections:
section B umbilical
WallD of Somite
Level of vesicle
umbilical Notochord

• Cresta neural à células que forman el sistema nervioso periférico y


section B Wall of E
vesicle
Primitive knot Primitive streak umbilical Intraembryonic coelom
Notochordal plate Intraembryonic F D
Primitive streak vesicle
Primitive knot B mesoderm
Notochordal plate Intraembryonic

autonómico.
Neural crest
Primitive knot Primitive streak
Cloacal membrane B
Notochordal plate Somites Intraembryonic
mesoderm
Caudal
A Cloacal membrane B Neural groovemesoderm neuropore

A Cloacal membrane Neural groove Neural crest


A Rostral Neural groove
Neural fold neuropore Neural crest
Rostral Neural crest
Neural fold Rostral
neuropore
Neural
Neural fold C
neuropore E Notochord
groove
Neural Levels of
Neural
groove sections:
Neural plate Levels Neural tube
groove
Levels of
D of Somite
Surface ectoderm
sections: Notochord Neural crest
Amnion sections:
Neural Neural D DE Somite Somite Notochord
ess groove fold Intraembryonic coelom
Notochord
EF D
Notochord
E
Intraembryonic
Intraembryonic coelom coelom
F F D D Neural crest
Somites Caudal
Wall of neuropore
Neural crest
Neural crest
Dorsal aorta
umbilical Somites
Somites Caudal
vesicle Caudal
neuropore
neuropore Umbilical vesicle
F
Notochordal plate Intraembryonic F I G U R E 1 7 – 1 The neural plate folds to form the neural tube. A, Dorsal view shows an embryo of approximately 17 days t
B mesoderm was exposed by removing the amnion. B, Transverse section of the embryo shows the neural plate and early development of the neu
groove and neural folds. C, Dorsal view of an embryo of approximately 22 days shows that the neural folds have fused opposite
C fourth to sixth somites but are spread apart at both ends. D to F, Transverse sections of the embryo at the levels shown in C illustr
Neural groove E Notochord
C formation of the neural tube and its detachment from the surface ectoderm. Some neuroectodermal cells are not included in the neu
Neural crest
C E Notochord
tube but remain between it and the surface ectoderm as the neural crest.
E Notochord
Neural tube
Surface ectoderm
Neuralcrest
Neural tube 18
Neural tube Surface ectoderm
Neural crest Notochord
Surface ectoderm
Neural crest Notochord
Notochord
Somite
Notochord
Intraembryonic coelom
D Dorsal aorta
Dorsal aorta
Neural crest Umbilical vesicle
Dorsal
F aorta Umbilical vesicle
F
F I G U R E 1 7 – 1 The neural plate folds to form the neural tube. A, Dorsal view shows an embryo of approximately 17 days that
380 THE DEVELOPING HUMAN

Oropharyngeal membrane
Neural plate

Neural plate Amnion


Neural Neural

Desarrollo del sistema nervioso


380 THE DEVELOPING HUMAN Notochordal process groove fold

Oropharyngeal membrane
Neural plate
Level of
Neural plate section B Wall of
Neural Neural Amnion
umbilical
Notochordal process groove fold vesicle

1. Neurulación à inicio de la formación de la placa y tubo neural à cuarta semana à en la


Primitive knot Primitive streak
Notochordal plate Intraembryonic
B mesoderm
región del 4to al 6to somita.
A
Level of
Cloacal membrane
section B
Neural groove
Wall of
umbilical
vesicle
Neural crest
Primitive Rostral
Primitive streak
Neural foldknot neuropore Notochordal plate Intraembryonic
B mesoderm
Neural Cloacal membrane
groove
A Levels of Neural groove
sections:
D Somite
Neural crest
Rostral Notochord
Neural fold E
neuropore
Intraembryonic coelom
F D
Neural
groove Neural crest
Levels of
Somites sections:
Caudal
neuropore
D Somite
Notochord
E
Intraembryonic coelom
F D
Neural crest
C Somites
Caudal E Notochord
neuropore

Neural tube
Surface ectoderm
Neural crest
Notochord
• Los dos tercios craneales de la placa y el tubo à Cerebro
C
• Tercio caudal de la placa y el tubo à Médula espinal E Notochord

Neural tube
Dorsal aorta Surface ectoderm 19
Neural crest
Umbilical vesicle Notochord
F
F I G U R E 1 7 – 1 The neural plate folds to form the neural tube. A, Dorsal view shows an embryo of approximately 17 days that
was exposed by removing the amnion. B, Transverse section of the embryo shows the neural plate and early development of the neural
groove and neural folds. C, Dorsal view of an embryo of approximately 22 days shows that the neural folds have fused opposite the
fourth to sixth somites but are spread apart at both ends. D to F, Transverse sections of the embryo at the levels shown in C illustrate
formation of the neural tube and its detachment from the surface
Dorsalectoderm.
aorta Some neuroectodermal cells are not included in the neural
tube but remain between it and the surface ectoderm as the neural crest.
Umbilical vesicle
F
382 THE DEVELOPING HUMAN

Rostral neuropore closing

Neural groove
Forebrain prominence

Heart prominence
Rostral neuropore

Desarrollo del sistema nervioso


Neural tube Omphaloenteric
Somites duct Somites

Connecting stalk
Caudal neuropore

A B
2. Fusión del pliegue neural y formación del tubo neural à comienza en el 5to somita y Caudal neuropore

continua en dirección craneal y caudal


Amnion
382 THE DEVELOPING HUMAN

Rostral neuropore closing


Rostral neuropore Otic pit
Amniotic cavity

Neural groove Pharyngeal arches


Forebrain prominence Notochord

Developing heart
Heart prominence
Rostral neuropore Neural tube

Lens placode
Umbilical Mesenchyme
Neural tube Omphaloenteric
Somites vesicle
Somites duct Omphaloenteric
duct
Neural canal
Connecting stalk Allantois
Caudal neuropore Umbilical cord

A B Upper limb bud


Caudal neuropore Connecting stalk D
Amnion C Caudal neuropore

F I G U R E 1 7 – 3 A, Dorsal view of an embryo of approximately 23 days shows fusion of the neural folds, which forms the neur
tube. B, Lateral view of an embryo of approximately 24 days shows the forebrain prominence and closing of the rostral neuropor
C, Diagrammatic sagittal section of the embryo at 23 days shows the transitory communication of the neural canal with the amniot
Rostral neuropore

• Lumen del tubo neural à canal neural (comunicación libre con la cavidad amniótica)
Otic pit cavity (arrows). D, In the lateral view of an embryo of approximately 27 days, notice that the neuropores shown in B are closed.
Amniotic cavity
Pharyngeal arches
Notochord
• El neuroporo rostral se cierra aproximadamenteDEVELOPMENT
Developing heart el día 25 OF SPINAL CORD Fig. 17-5 D). These neuroepithelial cells constitute th
ventricular zone (ependymal layer), which gives rise to a

• El neuroporo caudal se cierra aproximadamente of theel díaplate27and caudal eminence. The neural tube
The primordial spinal cord develops from the caudal part
Neural tube
neural
neurons and macroglial cells (macroglia) in the spina
cord (Fig. 17-6 ; see Fig. 17-5 E). Macroglial cells are th
caudal to the fourth pair of somites develops into the larger members of the neuroglial family of cells, whic
Lens placode
spinal cord (Fig. 17-5 ; see Figs. 17-3 and 17-4 ). The lateral
20
includes astrocytes and oligodendrocytes. Soon, a ma
Umbilical Mesenchyme
vesicle
walls of the neural tube thicken, gradually reducing the ginal zone composed of the outer parts of the neuroep
Omphaloenteric size of the neural canal until only a minute central canal thelial cells becomes recognizable (see Fig. 17-5 E). Th
duct of the spinal cord exists at 9 to 10 weeks (see Fig. 17-5 C). zone gradually becomes the white matter of the spina
Neural canal Retinoic acid signaling is essential in the development of cord as axons grow into it from nerve cell bodies in th
Allantois the spinal cord from early patterning to neurogenesis. spinal cord, spinal ganglia, and brain.
Umbilical cord
Initially, the wall of the neural tube is composed of a Some dividing neuroepithelial cells in the ventricula
Upper limb thick,
bud pseudostratified, columnar neuroepithelium (see zone differentiate into primordial neurons (neuroblasts
Connecting stalk D
C Caudal neuropore

F I G U R E 1 7 – 3 A, Dorsal view of an embryo of approximately 23 days shows fusion of the neural folds, which forms the neural
C H A P T E R 17 | NERVOUS SYSTEM 383

Midbrain flexure
Midbrain Hindbrain

Optic vesicle
Cervical flexure

Desarrollo del sistema nervioso Neural tube

Spinal ganglion
Somite

Notochord

3. Cierre de los neuroporos à establecimiento de circulación vascular para el tubo neural


Forebrain

Level of Aorta
section B

• Células neuroprogenitoras de la pared del tubo neural se engrosan à cerebro y médula


espinal Peritoneal cavity Midgut
C H A P T E R 17 | NERVOUS
A SYSTEM 383 B

Midbrain flexure Metencephalon


Midbrain Hindbrain
Pontine flexure

Optic vesicle
Mesencephalon
Cervical flexure
Myelencephalon

Neural tube Somite

Spinal ganglion Notochord Diencephalon


Optic cup

Primordial spinal cord

Forebrain Telencephalon

Level of Aorta
section B

Peritoneal cavity Midgut

A B C
F I G U R E 1 7 – 4 A, Schematic lateral view of an embryo of approximately 28 days shows the three primary brain vesicles: fore-
Metencephalon brain, midbrain, and hindbrain. Two flexures demarcate the primary divisions of the brain. B, Transverse section of the embryo shows
the neural tube that will develop into the spinal cord in this region. The spinal ganglia derived from the neural crest are also shown.
C, Schematic lateral view of the central nervous system of a 6-week embryo shows the secondary brain vesicles21
Pontine flexure and the pontine flexure
that occurs as the brain grows rapidly.

Mesencephalon

These embryonic
Myelencephalon cells form an intermediate zone (mantle migrate from the ventricular zone into the intermediate
layer) between the ventricular and marginal zones. Neu- and marginal zones. Some glioblasts become astroblasts
roblasts become neurons as they develop cytoplasmic and later astrocytes, whereas others become oligodendro-
Diencephalon processes (see Fig. 17-6 ). blasts and eventually oligodendrocytes (see Fig. 17-6 ).
Optic cup The supporting cells of the CNS, called glioblasts (spon- When the neuroepithelial cells cease producing neuro-
gioblasts), differentiate from neuroepithelial cells, mainly blasts and glioblasts, they differentiate into ependymal
Primordial
after spinal cord
neuroblast formation has ceased. The glioblasts cells, which form the ependyma (ependymal epithelium)
Telencephalon
C H A P T E R 17 | NERVOUS SYSTEM 389

Unfused neural Tuft of hair


arch Dura mater
Skin
Subarachnoid space
(containing CSF)

Spinal cord

Defectos en el desarrollo del sistema nervioso A


Back muscles

Vertebra
B

• NO cierre del tubo neural


Membranous sac

Dura mater
Open spinal cord

• Posiblemente 5 sitios de cierre involucrados en la formación del tubo neural Displaced spinal cord Skin

• Defectos por el NO cierre de cada sitio: Roots of spinal nerve

• Sitio 1: Espina bífida quística


Subarachnoid space

• Sitio 2: Meroencefalia (anencefalia)


Spinal ganglion

C D
F I G U R E 1 7 – 1 2 Diagrammatic sketches illustrate various types of spina bifida and the associated defects of the vertebral arches

• Sitio 2, 4 y 1: Craneoraquisquisis (one or more), spinal cord, and meninges. A, Spina bifida occulta. Observe the unfused neural arch. B, Spina bifida with meningocele.
C, Spina bifida with meningomyelocele. D, Spina bifida with myeloschisis. The defects illustrated in B to D are referred to collectively
as spina bifida cystica because of the cyst-like sac or cyst associated with them. CSF , Cerebrospinal fluid.

F I G U R E 1 7 – 1 3 A fetus at 20 weeks with severe neural tube defects,


including acrania, cerebral regression (meroencephaly), iniencephaly (enlarge-
ment of foramen magnum), and a sacral dimple (arrow).

22
Desarrollo de la médula espinal
• El tubo neural caudal al 4to par de somitas à Médula espinal
• Las paredes laterales se engrosan reduciendo gradualmente el tamaño del canal neural
384 THE DEVELOPING HUMAN

a las 9 a 10 semanas Neural canal


Marginal zone
Roof plate Dorsal median septum
Afferent neuroblasts Central canal
Primordium of in spinal ganglion Dorsal gray horn
Neural tube
spinal ganglion
Alar plate
Ventral
gray
horn
Sulcus
limitans
Motor
neuron
Basal plate

A
Ventral White
Motor neuroblast Floor plate
median matter
B Trunk of
fissure
spinal nerve
C Ventral motor root

Internal
limiting Dividing neuroepithelial cell
Mesenchyme
membrane
Spinal
External meninges
limiting
membrane

Ventricular zone Marginal zone


Neuroepithelial cells
Intermediate
D E (mantle) zone 23
F I G U R E 1 7 – 5 Development of the spinal cord. A, Transverse section of the neural tube of an embryo of approximately 23
days. B and C, Similar sections at 6 and 9 weeks, respectively. D, Section of the wall of the neural tube shown in A. E, Section of the
wall of the developing spinal cord shows its three zones. Notice that the neural canal of the neural tube is converted into the central
canal of the spinal cord (A to C).

lining the central canal of the spinal cord. SHH signal- Cell bodies in the alar plates form the dorsal gray
ing controls the proliferation, survival, and patterning columns, which extend the length of the spinal cord. In
of neuroepithelial progenitor cells by regulating GLI transverse sections of the cord, these columns are the
transcription factors (see Fig. 17-2 ). dorsal gray horns (see Fig. 17-7). Neurons in these
Microglia (microglial cells), which are scattered columns constitute afferent nuclei and groups of them
Roof plate Dorsal median septum
384 Neural
THE D E V E canal
LOPING HUMAN
Marginal zone Afferent neuroblasts Central canal
Primordium of in spinal ganglion Dorsal gray horn
Neural tube
spinal ganglion Roof plate Dorsal median septum
Neural canal Alar plate
Marginal zone Afferent Ventral
neuroblasts Central canal
Primordium of in spinal ganglion Dorsal gray horn
Neural tube gray
spinal ganglion
Alar plate horn
Sulcus Ventral
gray
limitans horn
Sulcus Motor
limitans neuron C H A P T E R Neuro
17
Basal plate (neuro
Motor Neural tube
neuron

Desarrollo de la médula espinal


Basal plate
A
A Ventral White
Motor neuroblast Floor plate
median matterVentral White
B FloorTrunk
plateof
B
Motor neuroblast
spinal nerveTrunk of
fissure median matter
fissure
C H A P T E R 17 | NERVOUS SYSTEM
spinal nerve
C Ventral motor root

• La pared del tubo neural está compuesta por un neuroepitelio:


Apolar neuroblast
Neuroepithelium Glioblas
C Ventral motor root
(neuroectoderm)
Neural tube
Internal
Internal
limiting
limiting Dividing Dividing
neuroepithelial cell
neuroepithelial cell Mesenchymal cell
Mesenchyme
Mesenchyme
membrane
membrane
C H A P T E R 17 | NERVOUS SYSTEM 385

Grueso External
External
limiting
Spinal
meninges
C H A P T E R 17
Spinal
meninges
NERVOUS SYSTEM | 385
Bipolar neuroblast
Neuroepithelium
(neuroectoderm)
Neuronas y células
Neural tubeC H| A P T Apolar
Pseudoestratificado
C H A P T E R 17 NERVOUS SYSTEM 385
limitingmembrane
macrogliales
Astroblas
membrane
E R 1 7neuroblast
NERVOUS | SYSTEM 385 Glioblast (spongioblast)
Microglial cell
Mesenchymal cell
en la médula espinal
Columnar Neuroepithelial cells
Ventricular zone Marginal zone
Mesenchymal cell
Unipolar neuroblast
Mesenchymal cell

Mesenchymal cell

Ventricular zone Marginal Neuroepithelium


T E R 1zone |
Neuroepithelium
CHA Intermediate
P 7 N E(neuroectoderm)
RVOUS SYSTEM 385
| C H A P T E R 17 NERVOUS SYSTEM 385

D Neuroepithelial cells (neuroectoderm)


E Neural tube Neural tube (mantle) zone
Intermediate ApolarNeuroepithelium
neuroblast Bipolar
Microglial cell neuroblast
Glioblast (spongioblast) Mesenchymal cell Ependyma
Neuroepithelium
F I G U R E 1 7 – 5 Development of the spinal cord. A, Transverse section Dendrite Microglial cell
(neuroectoderm) |
of the neural tube of an embryo of approximately 23
C H A P T E R 17 NERVOUS SYSTEM 385

D
days. B and C, Similar sections at 6 and 9 weeks,
E
respectively. D, Section
Neural (mantle)
of
tube
(neuroectoderm)
the wall
zone
of the neural tube shown in A. E, Section of the
Microglial cell Astroblast
Neural tube
Neuroepithelium
(neuroectoderm)

Oligodendroblast
Neural tube Mesenchymal cell
Microglial cell

F I GofU the
wall R E developing
1 7 – 5 Development
spinal cordofshows
the spinal cord. A,
its three Transverse
zones. Noticesection
that theof the neural
neural tubeofofthe
canal an neural
embryo tube of approximately
is converted 23 into the
Microglial cell central
Neuroepithelium
canal
days. Bofand
theC,spinal
Similarcord (A toatC).
sections 6 and 9 weeks, respectively. D, Section of the wall of the neural Glioblast
Apolar neuroblast tube shown
(spongioblast) Mesenchymal
in A. E, Section cell
of the Ependyma Neural tube
(neuroectoderm)
Apolar neuroblast Glioblast (spongioblast)
Microglial cell
Ependyma

wall of the developing spinal cord shows its three zones. Notice that the neural canal of the neural tube is converted into the central
C H A P T E R 17 | NERVOUS SYSTEM 385 Epithelium of

canal ofthethe spinal (A to C).


cordcanal
Bipolar neuroblast

lining central of the spinal cord. SHH signal- Apolar Cell bodies in theBipolar alar plates form the dorsalneuroblast
gray Ependyma
choroid plexus

Unipolar Protoplasmic astrocyte


Ependyma
Ependyma
Apolar neuroblast Glioblast (spongioblast)

Apolarneuroblast
neuroblast Glioblast (spongioblast)
Glioblast (spongioblast) Epithelium of
Astroblast Oligodendroblast

neuroblast
Neuroepithelium
ing controls the proliferation, survival, and patterning (neuroectoderm) columns, which extend the length of the spinal cord. In choroid plexus
| transverse
Unipolar neuroblast

of neuroepithelial progenitor cells


Neural
Apolar neuroblast
C H A P T E R Bipolar
tubeby regulating GLI
17 neuroblast NERVOUS SYSTEM sections of 385
Glioblast (spongioblast)
the cord, these columns
Bipolar neuroblast
Epithelium ofare the
Ependyma
choroid plexus
Dendrite
Astroblast Axon
Oligodendroblast
Epithelium of
choroid plexus

lining the central


transcription canal(see
factors of the
Fig.spinal
17-2 cord.
). SHH signal- Cell dorsal
bodies ingray the horns
alarAstroblast
plates
(seeform the dorsal
Fig.Oligodendroblast
17-7). graycell in these
Neurons
Microglial
Dendrite
Astroblast Oligodendroblast

Neuron
Oligodendrocyte

ingMicroglia
controls the(microglial
proliferation,cells),
survival,which are scattered
and patterning columns, columns
which extend
Mesenchymal constitute
cell the length afferent of thenuclei spinal and
Unipolar neuroblast cord.groups In of them
FEpithelium
Protoplasmic astrocyte Fibrous astrocyte

I G U RofE 1 7 – 6 Histogenesis of cells in the central nervous sy


throughout
of neuroepithelialthe gray and white
progenitor cellsmatter of the spinal
by regulating GLI cord, transverse form
Bipolar the dorsal
sections
neuroblast of thegray cord,columns. these columns As theare alarthe plates enlarge,Axon

Neuron

left) becomes a nerve cell or neuron. Neuroepithelial cells give rise t


Dendrite

are small cells that are derived from mesenchymal cells the dorsal median septum forms. Cell bodies in the basal
F I G U R E 1 7 – 6 Histogenesis of cells in the central nervous system. After further development, the multipolar neuroblast (lower
choroid plexus
(see Fig. invade
17-2 ). the CNS rather lateUnipolar dorsal gray horns
form (see
cell Fig. 17-7). Neurons in these
left) becomes a nerve cell or neuron. Neuroepithelial cells give rise to all neurons and macroglial cells. Microglial cells are derived from

transcription
(see Fig. 17-6factors). Microglia
neuroblast
in Bipolar
Mesenchymal
plates neuroblast the ventral Unipolar Astroblast
and lateral gray
neuroblast
mesenchymal cells that invade the developing nervous system with the blood

Oligodendroblast
columns. mesenchymal cells that Epithelium
invade of
vessels.
Oligodendrocyte

the developing nervous system with


theMicroglia (microglial it cells), whichpenetrated
are scattered columns constitute afferent nucleiofand thegroupsspinalof themthese columns choroid plexus
afterNeuroepithelium
Protoplasmic astrocyte Fibrous astrocyte

fetal period has been byDendrite


blood In transverse sections Axon cord,
fashion. Both processes of spinal ganglion cells have the
structural characteristics of axons, but the peripheral
the neural tube (primordium of the brain and spinal cord)
and form the primordial meninges (see Fig. 17-1F).
(neuroectoderm) Astroblast OligodendroblastProtoplasmic astrocyte
throughout the gray originate
and white matter of the spinal cord, form thearedorsal gray columns. As theand alar lateral
plates enlarge,
process is a dendrite in that there is conduction toward The external layer of these membranes thickens to

vessels. Microglia in the bone marrow and are the ventral gray horns gray horns,ofrespec- Fibrous astrocyte
Neuron

Neural tube Apolar Bipolar neuroblast


neuroblast Glioblast (spongioblast) F I G U R E 1 7 – 6 Histogenesis of cells
left) becomes a nerve cell or neuron.somatic
Epithelium
the cell body. The peripheral processes of spinal ganglion
Ependyma
cellspass in central
in the the spinal
or visceral
Neuroepithelial
nerves
nervous
structures
cells
to After
system.
(see
give rise to
sensory
allFig.
endings
further
17-8 ).
neurons and The
in
form the dura mater (Fig. 17-10 A and B), and the internal
layer,
development,
central cells.
macroglial
the pia arachnoid,
the multipolar
arachnoid
andMicroglial cells
is composed of pia mater
neuroblast (lower
mater (leptomeninges).
are derived from Fluid-filled spaces
Neuroepithelium
are small
part of the cells that are derived
mononuclear from
phagocytic mesenchymal
cell
(neuroectoderm)
population.cells the dorsaltively
Microglialmedian
Unipolar neuroblast
(see
cell septum
Fig. 17-5forms.
DendriteC). Cell
Axons bodies of in the
ventral basal
choroid
mesenchymal cells that invade the developing
horn
processes enter
nervous
plexus
cells
the system
Oligodendrocyte
roots of spinal nerves.
grow
spinal cord andblood
with the constitute
vessels.the dorsal appear within the leptomeninges that soon coalesce
to form the subarachnoid space (see Fig. 17-12A). The
origin of the pia mater and arachnoid from a single layer

Proliferation and differentiation the ventral roots Axonof the


invade the CNSofrather neuroepithelial out of
Astroblast
thethe spinal andcord and grayform
Oligodendroblast is indicated in the adult by arachnoid trabeculae, which

(see Fig.tube
17-6 ). Microglia late in plates form ventral lateral columns.
Development of Spinal Meninges
Neural are numerous, delicate strands of connective tissue that

Neural tube fashion. Both processes of spinal ganglion cells have the
The meninges (membranes covering the spinal cord) pass between the pia and arachnoid. Cerebrospinal
fashion. Both processes of spinal ganglion cells have the the neural tube (primordium of the brain and spinal cord)

cells in the developing spinal cord produce thick walls spinal


Microglialnerves.
cell As the basal
cord,platesastrocyte enlarge, they
these columns Neuronbulge
develop from cells of the neural crest and mesenchyme fluid (CSF) begins to form during the fifth week (see
structural characteristics of axons, but20 the
and peripheral and migrate
form the to primordial
surround meninges (see Fig. 17-1F).

the fetal period after it has been penetrated by blood Dendrite In transverse sections of the spinal Fibrous
between 35 days. The cells Fig. 17-12A).
Protoplasmic astrocyte process is a dendrite in that there is conduction toward The external layer of these membranes thickens to

and thin roof plates and floor plates (see Fig. 17-5 B).Axon ventrally
the cell body. The peripheral processes of spinal ganglion

on each side of the median plane. As this occurs,


cells pass in the spinal nerves to sensory endings in structural characteristics of axons, butOligodendrocyte
form the dura mater (Fig. 17-10 A and B), and the internal
layer, the pia arachnoid, is composed of pia mater the peripheral
vessels. Microglia originateofinthe the lateral
bone marrow andthe are spinal are the the ventral
Unipolar gray median
neuroblast horns andseptum lateral forms, gray horns, R Erespec-
somatic or visceral structures (see Fig. 17-8 ). The central and arachnoid mater (leptomeninges). Fluid-filled spaces

Differential thickening walls of ventral FIG Uand 1 7 – longitudinal


6 Histogenesis
processes enter the spinal cord and constitute the dorsal
roots of spinal nerves.
a deep process is aindendrite
of cells the central innervous
that there is conduction
system.
appear within the leptomeninges that soon coalesce
After further toward
to form the subarachnoid space (see Fig. 17-12A). The developme
part ofsoonthe mononuclear
produces aphagocytic cell population. tively (see Fig. 17-5 C). Axons of ventral horn cells grow
origin of the pia mater and arachnoid from a single layer

cord shallow longitudinal groove


Neuron
on groove (ventral median ofleft)
fissure)
Development Spinal becomes
develops
Meninges a nerve cell or neuron.
on the ventral the cell body. The peripheral
Neuroepithelial
Oligodendrocyte processes
cells give rise of spinal
to all neurons
is indicated in the adult by arachnoid trabeculae, which
are numerous, delicate strands of connective tissue that ganglion ce
and macroglial
Proliferation
side, the sulcusand Glioblast
differentiation
Unipolar
limitans of17-7;
F I G U R E 1 7 – 6 Histogenesis of cells in the central nervous system. After further development, the multipolar neuroblast (lower
neuroepithelial
neuroblast out ofneuron.
the spinal ofcord andgiveform
The meninges (membranes covering the spinal cord)

allthe ventral
mesenchymal roots ofC).
Epithelium the
cells ofthat
Protoplasmic cells
invadeastrocyte
the
pass between the pia and arachnoid. Cerebrospinal

from pass in theFibrous


developing nervous spinal nerves
system with the
astrocyte 24 vessels.
to blood
sensory endings in
Apolar neuroblast each (Fig. see
left)Fig. 17-5 surface the spinal cord (see andFig. 17-5
develop from cells of the neural crest and mesenchyme fluid (CSF) begins to form during the fifth week (see
Bipolar neuroblast
(spongioblast)
becomes B).
a nerve cell or Neuroepithelial
Ependyma cells rise to neurons macroglial cells. Microglial
between 20 and 35 days. The cells migrate to surround cells are derived Fig. 17-12A).

mesenchymal cellsDendrite choroid plexus somatic or visceral structures (see Fig. 17-8 ). The central
cells
This in the
groove developing
separatesspinal cord
the dorsal produce thick
part (alar walls
plate) fromspinal nerves. As the basal
Protoplasmicplates
that invade the developing nervous system with the blood vessels.
astrocyteenlarge, they bulge
Fibrous astrocyte
Apolar neuroblast
(basal
Glioblast (spongioblast) Astroblast Ependyma Oligodendroblast Axon processes enter the spinal cord and constitute the dorsal
the
and ventral
thin roofpart plates and plate).
Dendrite
The (see
floor plates alar Fig.
and17-5 basal B). plates ventrallyAxon on each side of the median
Development of Spinal plane. AsGanglia this occurs, roots of spinal nerves.
produce longitudinal bulges extending
Differential thickening of the lateral walls of the spinal through most of
the ventral median septum forms, and a deep
Neuron longitudinal Oligodendrocyte
the length of the developing spinal cord. fashion. This Both regional
processes of spinal
Neuron unipolar
The ganglion neurons
cells have the inneural
the the tube spinal ganglia
(primordium of the(dorsal rootcord)
brain and spinal
cord soon produces a shallow longitudinal structuralon
groove groove of(ventral
axons,F butI Gmedian
U R E fissure)
1 7 – 6develops the on
formfashion. the ventral
Both processes of the multipolar neuroblast (lower have the
spinal ganglion cells the neural tube (primordiu
characteristics the peripheral and Histogenesis primordial of cells
meninges in the
(see central
Fig. 17-1F). nervous system. After further development, the multipolar neurobl
separation is of fundamental importance because process is the
F I G U R E alar
a dendrite in thatganglia)
1 7 – 6 Histogenesis are
there left)
of cells
is conduction derived
toward from
in the central
Theneural
nervous crest
system.
external layer cells
After further
of these (Figs. 17-8 thickens
development,
Oligodendrocyte
membranes andDevelopment
to rise to all neurons of Spinal Meninges
each side, the
and basal plates are sulcus limitans (Fig. 17-7; see Fig.
later associated with afferentthe17-5
left)
cell B).
becomes
body. surface
aperipheral
nerve cell
and that 17-9
The of orthe spinal
neuron.
processes).theThe
of
becomescord
Neuroepithelial(see
axonsnervous
spinal ganglion
a nerveFig.
cells
ofProtoplasmic
cells
form 17-5
cell
givethe structural
or
rise
inwith C).
dura
neuron.
to
thethe all neurons
spinal
mater
astrocyte(Fig. characteristics
Neuroepithelial
and macroglial
ganglia are
17-10 A and B),
cells
cells. of giveaxons,
Microglial
at first
and
Fibrous the internal
astrocyte
but
cells are the
derived peripheral
from and macroglial and form
cells. the
Microglialprimordial
cells are der
Unipolar neuroblast cellsmesenchymal
pass in the cells invade tomesenchymal endings cells
developing that
system theprocess
invade pia the blood isunite
a dendrite
developing
vessels. in
nervous of that
piasystem
The there withis the
meninges conduction
blood towardcovering
vessels.
(membranes Thetheexternal
spinal layer
cord) of
This groove
Bipolar neuroblast efferent functions, the dorsal part (alar plate)
separatesrespectively. somatic from spinal nerves
or visceral structures bipolar,
Epithelium sensory
ofbut
(see Fig. 17-8 thecentral
). The
in layer,
two processes
and arachnoidsoon
arachnoid, is composed
mater (leptomeninges). in a Fluid-filled
T-shaped
mater
spaces
choroid plexus
Epithelium of the cell body. The peripheral processes
develop from spinalofganglion
of cells the neural formcrest theanddura mater (Fig.
mesenchyme
the ventral part (basal plate). The alar and basal
Bipolar neuroblast
plates
processes enter the spinal cordAxon and constitute the dorsal appear within the leptomeninges that soon coalesce

of Development of Spinal toorigin Ganglia


Astroblast roots ofProtoplasmic
Oligodendroblast spinal nerves. astrocyte choroid plexus Fibrous astrocyte form the cells pass inspace
subarachnoid the (see spinal
Fig. 17-12A). nerves
The to 20
between sensory
and 35endings
days. The in cellslayer, the to
migrate piasurround
arachnoid
produce longitudinal Dendritebulges extending
Astroblast through most Oligodendroblast Neuron of the pia mater and arachnoid from a single layer
somatic orbyvisceral and arachnoid mater (lep
is indicated in the adult arachnoid structures
trabeculae, which (see Fig. 17-8 ). The central
the length of the developing spinal Axoncord. ThisDevelopment
Fregional
fashion. Both Theof unipolar
processes
7 –Spinal
of neurons
spinal
Meninges in the
ganglion spinal
cells have ganglia
the
processesthe(dorsal
neural rootof(primordium
tube of the brain and spinal cord)
the dorsal
Istructural
G U R E characteristics
1 6 Histogenesis
of axons, of butcells
the in the central formenter
nervous system. the spinal
After cord
further andFig. constitute
development, the multipolar appear
neuroblast (lower within the lepto
are numerous, delicate strands connective tissue that
The meninges (membranes covering the spinal cord) peripheral and the primordial
pass between the pia and arachnoid. Cerebrospinal meninges (see 17-1F).
CH

Desarrollo de la médula espinal: neuronas Neural tube

• Algunas células neuroepiteliales en división en la zona ventricular à neuronas |


Neuroepithelium
C H A P T E R 17 NERVO

primordiales (neuroblastos). Neural tube


(neuroectoderm)

C H A P T E R 17 |
Apolar neuroblast
NERVOUS SYSTEM
Mesenc

• Estas células embrionarias forman una zona intermedia (capa de manto) entre
la las zonas ventricular y marginal. Apolar neuroblast Mesenchymal
Glioblast cell
(spongioblast)
Neural tube
Neuroepithelium
(neuroectoderm) Bipolar neuroblast

Microg

• Neuroblastos à neuronas Neuroepithelium


(neuroectoderm)
• A medida que desarrollan los procesos citoplasmáticos
Unipolar neuroblast
Neural tube
C H A P T E R 17 |
NERVOUS SYSTEM
Apolar neuroblast
385 Bipolar neuroblast
Glioblast (spongioblast)
Microglial cell
Epe
Dendrite
Astroblast O

Mesenchymal cell
Protoplasm

Apolar neuroblast Bipolar neuroblast Unipolar(spongioblast)


Glioblast neuroblast Axon Ependyma Epithe
Neuroepithelium choroid
(neuroectoderm) Astroblast Neuron
Oligodendroblast
Neural tube
Neural tube
Dendrite
25of cells in the cent
F I G U R E 1 7 – 6 Histogenesis
Microglial cell left) becomes a nerve cell or neuron. Neuroepithelial c
mesenchymal cells that invade the developing nervou

Unipolar neuroblast

Bipolar neuroblast Epithelium of


fashion.Protoplasmic
Both processes of spinal ganglionFibrou
astrocyte cells
Apolar neuroblast Glioblast (spongioblast) Ependyma choroid plexus
structural characteristics of axons, but the p
Dendrite
Astroblast Oligodendroblast
process is a dendrite in that there is conductio
Axon
|
(neuroectoderm)
Neural tube
Mesenchymal cell

Mesenchy

Neuroepithelium Mesenchy
(neuroectoderm)
Neural tube Neuroepithelium
Apolar neuroblast Glioblast (spongioblast)

|
(neuroectoderm)
Neural C H A P T E R 17
tube Microglial
NERV O U S Scell
YSTEM 38
Neuroepithelium Microglia

Desarrollo de la médula espinal: astrocitos y oligodentrocitos


(neuroectoderm)
Neural tube
Microglia
|
C H A P T E R 1 7Bipolar neuroblast
NERVOUS SYSTEM 385
• Glioblastos à Al cesar la formación de neuroblastos Mesenchymal cell
Apolar neuroblast Glioblast (spongioblast) Astroblast Epend
Oli
Apolar neuroblast Glioblast (spongioblast) Ependyma
• Glioblastos à Migran desde la zona ventricular a zonas intermedias y
marginales Neuroepithelium
Apolar neuroblast
(neuroectoderm) Glioblast (spongioblast)
Mesenchymal cell
Unipolar neuroblast Epend

• Glioblastos à Astrocitos o oligodentrocitos


Neural tube Bipolar neuroblast Epitheliu
choroid p
Microglial cell Dendrite
Astroblast Oligodendroblast

Bipolar neuroblast Epithelium of


Neuroepithelium choroid plexus
(neuroectoderm)
Neural tube Astroblast
Unipolar neuroblast Oligodendroblast
Protoplasmic astrocyte Fibrous a
Epitheliu
Bipolar neuroblast Microglial cell
C H A P T E R 17 | NERVOUS SYSTEM 385
Axon
choroid pl
Apolar neuroblast Glioblast (spongioblast)
Dendrite Astroblast Ependyma
Oligodendroblast
Neuron
F I G U R E 1 7 – 6 Histogenesis of cells in the central nervous system.Oligodend
After furth
Unipolar neuroblast left) becomes a nerve cell or neuron. Neuroepithelial cells give rise to all neurons and
Mesenchymal cellmesenchymal cells that invade the developing nervous system with the blood vesse
Protoplasmic astrocyte Fibrous astrocyte
Unipolar neuroblast
Dendrite
Apolar neuroblast Glioblast (spongioblast)
Axon Ependyma
Neuroepithelium
Bipolar neuroblast (neuroectoderm) fashion. Both processes of spinal ganglion
Neuron Epithelium of the
cells have the neural tub
Neural tube
Neural tube structural characteristics of choroid
axons, but plexus
the peripheral and form the
Dendrite F I G U R E 1 7 – 6 Microglial
Histogenesis of cells in the central nervous system. After further development,
cell process is a dendrite in that there is Oligodendrocyte
conduction toward
the
The
multipo
extern
left) becomes
Astroblasta nerve cell or neuron. Neuroepithelial
Oligodendroblast cells give rise to all neurons and macroglial cells. Microglial c
the cell body.
mesenchymal cells that invade the developing The peripheral
nervous system with processes
the blood spinal ganglion
of vessels. form the dura
cells pass in the spinal nerves to sensory endings in26 layer, the pi
somatic or visceral structures (see Fig. 17-8 ). The central and arachnoi
Oligodendr
Protoplasmic astrocyte processes enter the
Fibrous astrocyte spinal cord and constitute the dorsal appear withi
roots of spinal nerves. to form the s
fashion. Both processes of spinal ganglion cells have the Epithelium the neuraloftube (primordium of the
origin brap
of the
Bipolar neuroblast Glioblast
Apolar neuroblast
Unipolar neuroblast Axon (spongioblast) structural characteristics
Ependymaof axons, but the peripheral and form the primordial meninges
is indicated (see
in
Protoplasmic astrocyte
Development of Spinal Fibrous
choroid astrocyte
plexus
Meninges
process is a dendrite in that there is conduction toward The external layer of are these memb
numerous
Neuron theAstroblast
cell body. The peripheral processes of spinal(membranes
Oligodendroblast
The meninges ganglion covering form the thedura mater
spinal (Fig. 17-10
cord) pass A and B
between
cellsAxon
pass in the spinal nervesdevelop to sensory endings
from cells of theinneurallayer,
crest the pia arachnoid,fluid
and mesenchyme is compos
(CSF) b
F I G U R E 1 7 – 6 Histogenesis of cellssomatic
in the central nervous
or visceral system.
structures (see After further
Fig. 17-8 ). Thedevelopment,
central andthe multipolar
arachnoid neuroblast
mater (lower
(leptomeninges).
C H A P T E R 17
Neuroepithelium | NERVOUS SYSTEM 385
(neuroectoderm)
Neural tube
Microglial cell

Desarrollo de la médula espinal: células ependimales


Mesenchymal cell

• Células ependimales à Al cesar la producción de neuroblastos y glioblastos


Neuroepithelium
• Ependima à recubre el canal central de la médula espinal
(neuroectoderm)
Apolar
eural tube neuroblast Glioblast (spongioblast)
C H A P T E R 17 NERVOUS | Ependyma
SYSTEM 385
Microglial cell

Mesenchymal cell

Bipolar neuroblast
oblast Glioblast (spongioblast) Ependyma Epithelium of
Neuroepithelium choroid plexus
(neuroectoderm)
Neural tube
Neural tube Astroblast Oligodendroblast
Microglial cell

roblast Epithelium of
Unipolar neuroblast choroid plexus 27
Apolar neuroblast Astroblast Glioblast (spongioblast)
Oligodendroblast Ependyma

drite

roblast Oligodendrocyte
Desarrollo de la médula espinal: células microgliales
C H A P T E R 17 | NERVOUS SYSTEM 385
• Microglia = células microgliales (dispersas por la materia gris y blanca)
• Células pequeñas que se derivan de las células mesenquimales
C H A •P Se
T Eoriginan
• Invade el SNC
|
R 17 en la médula
NERVOUS ósea SYSTEM 385
Mesenchymal cell

Neuroepithelium
(neuroectoderm)
tube
Mesenchymal cell Microglial cell

st Glioblast (spongioblast) Ependyma 28


Microglial cell
Desarrollo de la médula espinal
• Proliferación y diferenciación de células neuroepiteliales à paredes gruesas y placas
G HUMAN

386 THE DEVELOPING HUMAN


Roof plate Dorsal median septum
Marginal zone Afferent neuroblasts Central canal Roof plate
mordium of in spinal ganglion Dorsal gray horn
nal ganglion Dorsal root of Alar plate
Alar plate spinal nerve
Ventral
F I G U R E 1 7 – 7 Transverse section of an embryo (×100) at Sulcus limitans
gray
Carnegie stage 16 at approximately 40 days. The ventral root of Central canal
horn
the spinal nerve is composed of nerve fibers arising from neuro- Neuroepithelium
Sulcus
blasts in the basal plate (developing ventral horn of spinal cord),
limitans Spinal ganglion
whereas the dorsal root is formed by nerve processes arising from Basal plate
Motor
neuroblasts in the spinal ganglion. (From Moore KL, Persaud TVN,
neuron Floor plate
Shiota K: Color
Basal plate atlas of clinical embryology, ed 2, Philadelphia,
2000, Saunders.)
Developing
body of vertebra
Ventral root of
Ventral White
Motor neuroblast Floor plate spinal nerve
median matter 29
B Trunk of
fissure
spinal nerve
C Ventral motor root

Neural crest
Dividing neuroepithelial cell
6 Desarrollo de la médula espinal
THE DEVELOPING HUMAN

Roof plate

Dorsal root of Alar plate


spinal nerve
• Los cuerpos celulares en las placas alares
G U R E 1 7 – 7 Transverse section of an embryo (×100) at Sulcus limitans
forman
rnegie stage columnas 40 dorsales
16 at approximately querootseof
days. The ventral Central canal
extienden
spinal nerve a lo largo
is composed defibers
of nerve la médula espinal
arising from neuro- Neuroepithelium
sts in the basal plate (developing ventral horn of spinal cord),
ereas the dorsal root is formed by nerve processes arising from Spinal ganglion Basal plate
• Las
uroblasts neuronas
in the spinal en Moore
ganglion. (From estasKL, Persaud
columnasTVN,
Floor plate
constituyen
ota K: Color núcleos
atlas of clinical aferentes
embryology, y grupos
ed 2, Philadelphia,
00, Saunders.)
de ellas forman columnas grises Developing
dorsales. body of vertebra
Ventral root of
spinal nerve

30

Neural crest

Neural crest cells Neural crest cells


Desarrollo de la médula espinal
DEVELOPING HUMAN

Roof plate Dorsal median septum


Neural canal
• Cuernos grises ventrales
Marginal zone Afferent neuroblasts Central canal
Primordium of Dorsal gray horn
be • Cuernos grises laterales
spinal ganglion
in spinal ganglion
Alar plate
Ventral
• Axones à crecen fuera de la médula gray
espinal à forman raíces ventrales de horn
Sulcus
limitans
• Las placas basales se agranda, se hinchan Motor
de los lados para formar el tabique
Basal plate
neuron
medianoventral y se desarrolla el surco
longitudinal profundo.
Ventral White
Motor neuroblast Floor plate
median matter
B Trunk of
fissure
spinal nerve
C Ventral motor root
31

Dividing neuroepithelial cell


Mesenchyme
ne
Spinal
meninges
Roof plate

Dorsal root of Alar plate


spinal nerve
F I G U R E 1 7 – 7 Transverse section of an embryo (×100) at Sulcus limitans
Carnegie stage 16 at approximately 40 days. The ventral root of Central canal
the spinal nerve is composed of nerve fibers arising from neuro- Neuroepithelium
blasts in the basal plate (developing ventral horn of spinal cord),
whereas the dorsal root is formed by nerve processes arising from Spinal ganglion Basal plate
neuroblasts in the spinal ganglion. (From Moore KL, Persaud TVN,
Floor plate
Shiota K: Color atlas of clinical embryology, ed 2, Philadelphia,
2000, Saunders.)
Developing

Desarrollo de la médula espinal: ganglios espinales


body of vertebra
Ventral root of
spinal nerve

Neural crest CHA


• Células de la cresta neural à neuronas
of the first lum
Neural crest cells Neural crest cells unipolares
Positional en ganglios
Changes espinales.
of Spinal Cord average level b
Neural tube The spinal cord in the embryo extends the entire length adults may be
of the vertebral canal (see Fig. 17-10 A). The spinal nerves or as inferior
Dorsal gray
pass through the intervertebral foramina opposite their nerve roots, e
Spinal ganglion
horn Spinal cord Dorsal levels of origin. Because the vertebral column and dura segments, run
root
Site of Unipolar neuron mater grow more rapidly than the spinal cord, this posi- responding lev
(spinal ganglion cell)
lateral gray
horn
tional relationship of the spinal nerves does not persist. The nerve roo
The caudal end of the spinal cord in fetuses gradually cone) form a b
Ventral Satellite cell comes to lie at relatively higher levels. In a 24-week-old equina (Latin
gray
horn fetus, it lies at the level of the first sacral vertebra (see cral enlargem
Spinal nerve
Ventral root
Schwann cell Fig. 17-10 B). spinal cord (se
White communicating ramus
The spinal cord in neonates terminates at the level of Although th
Multipolar neuron the second or third lumbar vertebra (see Fig. 17-10 C). In end at the S2
(sympathetic ganglion cell) adults, the cord usually terminates at the inferior border Distal to the c
forms a long fi
Ganglion of sympathetic trunk
filum), which i
Melanocyte of the embryo
(Latin thread)
attaches to the
Suprarenal medulla (see Fig. 17-10
(chromaffin cells) A B C D
Celiac Renal Suprarenal gland
Neural Bipolar Unipolar
ganglion ganglion
crest cell neuroblasts afferent Myelinatio
neuron
32 Myelin sheath
Plexus in
intestinal tract
F I G U R E 1 7 – 8 Diagram shows some derivatives (arrows) of the neural crest. Neural crest cells also differentiate into the cells
F I G U R E 1 7 – 9 A to D, Diagrams show successive stages cord begin to
in the afferent ganglia of cranial nerves and many other structures (see Chapter 5, Fig. 5-5). Formation of a spinal nerve is also in the differentiation of a neural crest cell into a unipolar afferent continue to f
illustrated. neuron in a spinal ganglion. Arrows indicate how a unipolar 17-11E). Mye
neuron is formed. peptide isoform

Body of vertebra Spinal cord


tional relationship of the spinal nerves does not persist. The nerve roots inferior to the end of the cord (medullary
The caudal end of the spinal cord in fetuses gradually cone) form a bundle of spinal nerve roots called the cauda
comes to lie at relatively higher levels. In a 24-week-old equina (Latin horse tail), which arises from the lumbosa-
fetus, it lies at the level of the first sacral vertebra (see cral enlargement (swelling) and medullary cone of the
Fig. 17-10 B). spinal cord (see Fig. 17-10 D).
The spinal cord in neonates terminates at the level of Although the dura mater and arachnoid mater usually
the second or third lumbar vertebra (see Fig. 17-10 C). In end at the S2 vertebra in adults, the pia mater does not.
adults, the cord usually terminates at the inferior border Distal to the caudal end of the spinal cord, the pia mater
forms a long fibrous thread, the filum terminale (terminal
filum), which indicates the original level of the caudal end
of the embryonic spinal cord (see Fig. 17-10 C). The filum

Desarrollo de la médula espinal: meninges espinales


A B C D
(Latin thread) extends from the medullary cone and
attaches to the periosteum of the first coccygeal vertebra
(see Fig. 17-10 D).

Neural Bipolar Unipolar


Myelination of Nerve Fibers
• Meninges: membranas que cubren
crest cell neuroblasts
laaround
Myelin sheaths médula espinal
afferent
neuron
the nerve fibers within the spinal
FIGURE 17–9 cord begin to form during the late fetal period and
• Se desarrollan a partir de las células deduring
la cresta neural y el mesénquima
A to D, Diagrams show successive stages
continue to form the first postnatal
in the differentiation of a neural crest cell into a unipolar afferent year (Fig.
17-11E). Myelin basic proteins, a family of related poly-
neuron in a spinal ganglion. Arrows indicate how a unipolar
• Entre los 20 y 35 días
neuron is formed. peptide isoforms, are essential in myelination; β -integrins 1

Body of vertebra Spinal cord Spinal cord

• Engrosamiento de capa externa à duramadre


Spinal cord
y capa interna (pia aracnoidea)
Lumbosacral
L1 L1 L1 L1
enlargement
Dura mater
Pia mater
• Aparecen espacios llenos de líquido dentro de
Medullary
cone
Intervertebral
foramen Arachnoid las leptomeninges que se unen à espacio L3

subaracnoideo.
S1
Root of
1st sacral Filum terminale
nerve S1
Medullary cone S1

Spinal ganglion Root of


C1 1st sacral
nerve S1

C1

C1
End of dural sac 33

C1

A B C D

Attachment of
filum terminale
F I G U R E 1 7 – 1 0 Diagrams show the position of the caudal end of the spinal cord in relation to the vertebral column and
fetus, it lies at the level of the first sacral vertebra (see cral enlargement (swelling) and medullary cone of the
Fig. 17-10 B). spinal cord (see Fig. 17-10 D).
The spinal cord in neonates terminates at the level of Although the dura mater and arachnoid mater usually
the second or third lumbar vertebra (see Fig. 17-10 C). In end at the S2 vertebra in adults, the pia mater does not.
adults, the cord usually terminates at the inferior border Distal to the caudal end of the spinal cord, the pia mater
forms a long fibrous thread, the filum terminale (terminal
filum), which indicates the original level of the caudal end
of the embryonic spinal cord (see Fig. 17-10 C). The filum
(Latin thread) extends from the medullary cone and
attaches to the periosteum of the first coccygeal vertebra
(see Fig. 17-10 D).
A B C D

Desarrollo de la médula espinal: Cambios posicionales


Neural Bipolar Unipolar
crest cell neuroblasts afferent Myelination of Nerve Fibers
neuron Myelin sheaths around the nerve fibers within the spinal
F I G U R E 1 7 – 9 A to D, Diagrams show successive stages cord begin to form during the late fetal period and
in the differentiation of a neural crest cell into a unipolar afferent continue to form during the first postnatal year (Fig.
MÉDULA ESPINAL EMBRIÓN neuron in a spinal ganglion. Arrows indicate how a unipolar MÉDULA ESPINAL RECIEN NACIDO
17-11E). Myelin basic proteins, a family of related poly-
neuron is formed. peptide isoforms, are essential in myelination; β1-integrins
Extendida a lo largo del canal vértebral Termina al nivel de la 2da o 3era vértebra lumbar
Body of vertebra Spinal cord Spinal cord

Spinal cord Lumbosacral


L1 L1 L1 L1
enlargement
Dura mater
Pia mater
Medullary
cone
Intervertebral
foramen Arachnoid L3
S1
Root of
1st sacral Filum terminale
nerve S1
Medullary cone S1

Spinal ganglion Root of


C1 1st sacral
nerve S1

C1

C1
End of dural sac

C1

A B C D

MÉDULA ESPINAL FETO 24 SEMANAS Attachment of

MÉDULA ESPINAL ADULTOS


filum terminale

Se encuentra al nivel de la 1era vértebra


F I G U R E 1 7 – 1 0 Diagrams show the position of the caudal end of the spinal cord in relation to the vertebral column and
34
Termina en el borde inferior de la primera
meninges at various stages of development. The increasing inclination of the root of the first sacral nerve is also illustrated. A, At 8

sacra weeks. B, At 24 weeks. C, Neonate. D, Adult.


vértebra lumbar
Desarrollo de la médula espinal: mielinización de las
fibras nerviosas
• Formación de vainas de mielina alrededor de las fibras nerviosas dentro de la médula
espinal
• Período fetal tardío hasta el 1er año postnatal
• Formadas por oligodendrocitos
• Se envuelven alrededor del axón formando varias capas

35
Membranous sac

Dura mater
Open spinal cord

Displaced spinal cord

Roots of spinal nerve

Subarachnoid space

Spinal ganglion

C D

Defectos de nacimiento de la médula espinal


F I G U R E 1 7 – 1 2 Diagrammatic sketches illustrate various types of spina bifida and the associated defects of the
(one or more), spinal cord, and meninges. A, Spina bifida occulta. Observe the unfused neural arch. B, Spina bifida wit
C, Spina bifida with meningomyelocele. D, Spina bifida with myeloschisis. The defects illustrated in B to D are referre
as spina bifida cystica because of the cyst-like sac or cyst associated with them. CSF , Cerebrospinal fluid.

• La mayoría resultan del fracaso de la fusión de uno o


más arcos neurales de las vértebras en desarrollo
durante la cuarta semana
F I G U R E 1 7 – 1 3 A fetus at 20 weeks with severe neur
including acrania, cerebral regression (meroencephaly), inienc

• Afectan los tejidos que recubren la médula espinal:


ment of foramen magnum), and a sacral dimple (arrow).

• Meninges
• Arcos neurales
• Músculos
• Piel

• Defectos que involucran los arcos neurales: ESPINA


BÍFIDA
• Subtipos que se basan en el grado y patrón del defecto

• Espina bífida = NO fusión de las mitades de los arcos


neurales embrionarios 36
C H A P T E R 17 | NERVOUS SYSTEM 389

Unfused neural Tuft of hair


arch Dura mater
Skin
Subarachnoid space
(containing CSF)

Spinal cord

Back muscles

Vertebra
A B
ESPINA BÍFIDA OCULTA ESPINA BÍFIDA CON MENINGOCELE
Membranous sac

Dura mater
Open spinal cord

Displaced spinal cord Skin

Roots of spinal nerve

Subarachnoid space

Spinal ganglion

C D 37

ESPINA BÍFIDA CON MENINGOMIELOCELE ESPINA BÍFIDA CON MIELOSQUISIS


F I G U R E 1 7 – 1 2 Diagrammatic sketches illustrate various types of spina bifida and the associated defects of the vertebral arches
(one or more), spinal cord, and meninges. A, Spina bifida occulta. Observe the unfused neural arch. B, Spina bifida with meningocele.
C, Spina bifida with meningomyelocele. D, Spina bifida with myeloschisis. The defects illustrated in B to D are referred to collectively
as spina bifida cystica because of the cyst-like sac or cyst associated with them. CSF , Cerebrospinal fluid.
S

C
F I G U R E 1 7 – 1 2 Diagrammatic sketches illustrate va

Defectos de nacimiento de la médula espinal: seno


(one or more), spinal cord, and meninges. A, Spina bifida o
C, Spina bifida with meningomyelocele. D, Spina bifida wit
as spina bifida cystica because of the cyst-like sac or cyst a

dérmico
• Invaginación cutánea revestida de epidermis
• Se extiende desde la piel hasta la médula espinal
• Asociado con el cierre del tubo neural y la formación
de meninges
• Causa: falla del ectodermo superficial para
desprenderse del neuroectodermo y las meninges que F
lo envuelven inc
m
• Resultado: Meninges continuas con un canal estrecho
que se extiende hasta un hoyuelo en la piel de la
región sacra de la espalda.

38
Defectos de nacimiento de la médula espinal: espina
bífida oculta
• Defecto en el tubo neural
• Causa: falla de la fusión de las mitades de uno o más
arcos neurales para fusionarse en el plano medio
• Ocurre en la vértebra L5 o S1
• Síntomas (generalmente NO produce)
• Pequeño hoyuelo con un mechón de cabello que surge de él
• Lipoma suprarrenal
• Seno dérmico
• Otras marcas de nacimiento C H A P T E R 17 | NERVOUS SYSTEM

Unfused neural Tuft of hair


arch Dura mater
Skin
Subarachnoid space
(containing CSF)

Spinal cord

Back muscles

Vertebra
A B 39

Membranous sac

Dura mater
Open spinal cord

Displaced spinal cord Skin

Roots of spinal nerve


Defectos de nacimiento de la médula espinal: espina
bífida quística
• Tipos severos de espína bífida
• Implican protrusión de la médula espinal y/o
meninges
• Defectos de arcos vertebrales
• 1/5000 nacimientos
• Tipos:
• Meningocele: quiste con meninges y LCR = defecto en
columna vertebral
• Meningomomielo: médula o raíces nerviosas contenidas C H A P T E R 17 | NERVOUS SYSTEM 389

dentro del quiste Unfused neural Tuft of hair


arch Dura mater

• Involucran ausencia de la mayor parte del cerebro,


Skin
Subarachnoid space

anomalías faciales, parálisis parcial o completa del


(containing CSF)

músculo esquelético entre otras.


Spinal cord

Back muscles

A
Vertebra
B 40

Membranous sac

Dura mater
Open spinal cord

Displaced spinal cord Skin

Roots of spinal nerve


neuropore. (Modified from Jones KL: Smith’s recognizable patterns of human malformations, ed 4, Philadelphia, 1988, Saunders.)
brain) results from defective closure of the rostral neuropore, and meningomyelocele results from defective closure of the caudal
F I G U R E 1 7 – 1 7 Schematic illustration shows the embryologic basis of neural tube defects. Meroencephaly (partial absence of

Meroencephaly
Defectos de nacimiento de la médula espinal:

Clubfoot

Myeloschisis
meningomielocele

caudal to lesion
+/– Clubfoot

Neural deficit
• Más común y más grave que la espina bífida con
meningocele
• Ocurre en cualquier lugar de la columna vertebral

Meningomyelocele
+/– Hydrocephalus
|
(más comunes en la región lumbar y sacra)
C H A P T E R 17 NERVOUS SYSTEM 389

• Hidrocefalia asociada (coexistencia de una


Unfused neural Tuft of hair
arch Dura mater
Skin
Subarachnoid space

malformación de Arnold-Chiari) (containing CSF)

Skin
• Asociado con craneo-lacunia (desarrollo defectuoso

Spina bifida occulta


Spinal cord

Unfused vertebral
Tuft of hair

arch
de la calvaria) à áreas deprimidas y no densificadas
Back muscles

Vertebra
A B

en las superficies internas de los huesos

Dura mater
Membranous sac

Vertebra

Spinal cord

vertebral arch
Incomplete

Subarachnoid space
Dura mater
Open spinal cord

Displaced spinal cord Skin

Roots of spinal nerve

Subarachnoid space

Spinal ganglion

C D
41
F I G U R E 1 7 – 1 2 Diagrammatic sketches illustrate various types of spina bifida and the associated defects of the vertebral arches
(one or more), spinal cord, and meninges. A, Spina bifida occulta. Observe the unfused neural arch. B, Spina bifida with meningocele.
C, Spina bifida with meningomyelocele. D, Spina bifida with myeloschisis. The defects illustrated in B to D are referred to collectively
as spina bifida cystica because of the cyst-like sac or cyst associated with them. CSF , Cerebrospinal fluid.
Defectos de nacimiento de la médula espinal:
mielosquisis
• Tipo más severo de espina bífida
|
• La médula espinal esta abierta en el área afectada, C H A P T E R 17 NERVOUS SYSTEM 389

porque los pliegues neurales NO se fusionaron.


Unfused neural
arch
Tuft of hair
Dura mater

• Resultado: médula espinal está representada por una


Skin
Subarachnoid space
(containing CSF)

masa aplanada de tejido nervioso Spinal cord

• Síntomas: parálisis permanente o debilidad en Back muscles

extremidades
A
inferiores B
Vertebra

Membranous sac

Dura mater
Open spinal cord

Displaced spinal cord Skin

Roots of spinal nerve

Subarachnoid space

Spinal ganglion

C D 42
F I G U R E 1 7 – 1 2 Diagrammatic sketches illustrate various types of spina bifida and the associated defects of the vertebral arches
(one or more), spinal cord, and meninges. A, Spina bifida occulta. Observe the unfused neural arch. B, Spina bifida with meningocele.
C, Spina bifida with meningomyelocele. D, Spina bifida with myeloschisis. The defects illustrated in B to D are referred to collectively
as spina bifida cystica because of the cyst-like sac or cyst associated with them. CSF , Cerebrospinal fluid.
Defectos de nacimiento de la médula espinal: CAUSAS
• Factores nutricionales y ambientales
• Interacciones genéticas y epigenéticas

• La fortificación con ácido fólico y suplementos de ácido fólico tomados antes de la


concepción y continuados durante al menos 3 meses reducen la incidencia
• Bajos niveles de B12 en la madre podrían aumentar significativamente el riesgo

43
Desarrollo del cerebro
• Comienza durante la 3era semana
• Cuando la placa y el tubo neural se desarrollan a partir del neuroectodermo

• Tubo neural craneal al 4to par de somitas à Cerebro

• Células neuroprogenitoras migran y se diferencian para formar áreas específicas del


cerebro

• Fusión de los pliegues neurales en la región craneal + cierre de neuroporo = 3 vesículas


cerebrales a partir de las cuales se desarrolla el cerebro:
1. Cerebro anterior (proencéfalo)
2. Cerebro medio (mesencéfalo)
3. Rombencéfalo

44
Defectos de nacimiento en el desarrollo del cerebro
• Debido a la complejidad de su historia embriológica, es común el desarrollo anormal del
cerebro
• 3 de cada 1000 nacimientos
• La mayoría de los defectos congénitos son el resultado del cierre defectuoso del neuroporo
rostral durante la 4ta semana
• Involucran los tejidos suprayacentes (meninges y calvaria).
• Causa: Factores genéticos, nutricionales y ambientales
• Pueden ser alteraciones en:
• La morfogénesis
• La histogénesis del tejido nervioso à convulsiones y varios grados de retraso mental
• Fallas del desarrollo (notocorda, somitas, mesémquima y cráneo

45
Defectos de nacimiento en el desarrollo del cerebro:
hipófisis faríngea y craneofaringioma
• Se forma al persistir un remanente del tallo del
divertículo hipofisario
• Forma una hipófisis faríngea en el techo de la
orofarínge

• Algunas veces se forman masas del tejido del lóbulo


anterior fuera de la cápsula de la glándula pituitaria,
dentro de la silla turca del hueso esfenoides
• Ocasionalmente, se desarrolla un tumor benigno
(craneofaringioma) en la silla turca
• Con menos frecuencia pueden desarrollarse en la
faringe o la base fenoide
46
|

Central sulcus
Lateral sulcus
Insula

Defectos de nacimiento en el desarrollo del cerebro:


Frontal pole Occipital pole
Cerebellum
Pons B C
A Medulla oblongata

Encefalocele
F I G U R E 1 7 – 3 0 A, Lateral view of the brain of a fetus that died before delivery (25 weeks). B, The medial (top) and la
(bottom) surfaces of the fetal brain (week 25). C, The lateral (top) and medial (bottom) surfaces of the fetal brain at week 38 (labe
photo: 40 weeks from last normal menstrual period). As the brain enlarges, the gyral pattern of the cerebral hemispheres beco
more complex (compare with Figure 17-29 ). (A, From Nishimura H, Semba R, Tanimura T, Tanaka O: Prenatal development o
human with special reference to craniofacial structures: an atlas, Bethesda, MD, 1977, U.S. Department of Health, Education,
Welfare, National Institutes of Health.)

• Hernia del contenido intracraneal


• Resultante de un defecto en el cráneo Neurocranium
Defect in cranium at foramen magnum

(cráneo bífido) Skin

• Comunes en la región occipital


• Ocurre en 1 de 2000 nacimientos aprox. Part of cerebellum

A C

• La hernia puede contener:


Occipital lobe Defect at posterior fontanelle of cranium

• Meninges (meningocele) Arachnoid mater


Ventricle Dura mater
Subarachnoid
• Meninges y parte del cerebro space
(dilated)

(meningoencefalocele) Dura mater


Part of occipital lobe

• Meninges, parte del cerebro y parte del sistema B D


Defect at posterior fontanelle of cranium
Skin

ventricular (menngohidroencefalocele) F I G U R E 1 7 – 3 1 Schematic drawings illustrate encephalocele (cranium bifidum) and various types of herniation of the brain
meninges. A, Sketch of the head of a neonate with a large protrusion from the occipital region of the cranium. The upper red c
indicates a cranial defect at the posterior fontanelle (membranous interval between cranial bones). The lower red circle indicat
cranial defect near the foramen magnum. B, Meningocele consists of a protrusion of the cranial meninges that is filled with cere
spinal fluid. C, Meningoencephalocele consists of a protrusion of part of the cerebellum that is covered by meninges and
D, Meningohydroencephalocele consists of a protrusion of part of the occipital lobe that contains part of the posterior horn of a la
ventricle. 47
e Occipital pole
Cerebellum
ns B C
A Medulla oblongata
1 7 – 3 0 A, Lateral view of the brain of a fetus that died before delivery (25 weeks). B, The medial (top) and lateral
faces of the fetal brain (week 25). C, The lateral (top) and medial (bottom) surfaces of the fetal brain at week 38 (label on
eeks from last normal menstrual period). As the brain enlarges, the gyral pattern of the cerebral hemispheres becomes
ex (compare with Figure 17-29 ). (A, From Nishimura H, Semba R, Tanimura T, Tanaka O: Prenatal development of the
special reference to craniofacial structures: an atlas, Bethesda, MD, 1977, U.S. Department of Health, Education, and
tional Institutes of Health.)

Neurocranium
Defect in cranium at foramen magnum

Skin

Part of cerebellum

C
Occipital lobe Defect at posterior fontanelle of cranium C H A P T E R 17 | NERVOUS SYSTEM 407

Ventricle Dura mater


Subarachnoid Arachnoid mater
space
(dilated)
Part of occipital lobe
Dura mater

Skin
D
Defect at posterior fontanelle of cranium
1 7 – 3 1 Schematic drawings illustrate encephalocele (cranium bifidum) and various types of herniation of the brain and
A, Sketch of the head of a neonate with a large protrusion from the occipital region of the cranium. The upper red circle
cranial defect at the posterior fontanelle (membranous interval between cranial bones). The lower red circle indicates a
ct near the foramen magnum. B, Meningocele consists of a protrusion of the cranial meninges that is filled with cerebro-
C, Meningoencephalocele consists of a protrusion of part of the cerebellum that is covered by meninges and skin.
hydroencephalocele consists of a protrusion of part of the occipital lobe that contains part of the posterior horn of a lateral

A B
F I G U R E 1 7 – 3 4 Magnetic resonance images (MRIs) of a 1-day-old neonate show a meningocele. A, Sagittal MRI taken so that
the cerebrospinal fluid (CSF) appears bright. The image is blurred because of movement of the neonate. B, Axial image located at the
cranial defect near the foramen magnum and taken so that CSF appears dark. Compare with Figure 17-31C.
Meningoencefalocele en el área occipital Meningocele

48

F I G U R E 1 7 – 3 5 A, Sonogram of a normal fetus at 12 weeks (left)


and a fetus at 14 weeks with acrania and meroencephaly (right). B, Mag-
netic resonance image of diamniotic-monochorionic twins, one with mero-
encephaly. Notice the absent calvaria (white arrow) of the abnormal
twin and the amnion of the normal twin (black arrow). (A, From Pooh RK,
Pooh KH: Transvaginal 3D and Doppler ultrasonography of the fetal brain,
Semin Perinatol 25:38, 2001.)
C H A P T E R 17 | NERVOUS SYSTEM 407

degeneración
posterior y la calvaria
Meroencefalia

compatible con la vida


cerrarse en la 4ta semana

• 1 de cada 1000 nacimientos


A B
F I G U R E 1 7 – 3 4 Magnetic resonance images (MRIs) of a 1-day-old neonate show a meningocele. A, Sagittal MRI taken so that
the cerebrospinal fluid (CSF) appears bright. The image is blurred because of movement of the neonate. B, Axial image located at the

• Diagnostico: US, MRI, radiografía


cranial defect near the foramen magnum and taken so that CSF appears dark. Compare with Figure 17-31C.
• Defecto grave de la calvaria y el cerebro

extruido del cráneo (exenfalia) à


• Origen: falla del neuroporo rostral en

• La mayor parte del cerebro está expuesto

• Pueden sobrevivir brevemente pero no es


mesencéfalo, la mayoría del cerebro
• Resultado: ausencia del cerebro anterior, el

B
A

F I G U R E 1 7 – 3 5 A, Sonogram of a normal fetus at 12 weeks (left)


A

and a fetus at 14 weeks with acrania and meroencephaly (right). B, Mag-


netic resonance image of diamniotic-monochorionic twins, one with mero-
encephaly. Notice the absent calvaria (white arrow) of the abnormal
twin and the amnion of the normal twin (black arrow). (A, From Pooh RK,
Pooh KH: Transvaginal 3D and Doppler ultrasonography of the fetal brain,
Semin Perinatol 25:38, 2001.)
Defectos de nacimiento en el desarrollo del cerebro:

B
49
F I G U R E 1 7 – 3 4 Magnetic resonance images (MRIs) of a 1-day-old

and a fetus at
netic resonanc

Pooh KH: Tran


the cerebrospinal fluid (CSF) appears bright. The image is blurred because

twin and the a


FIGURE 1

Semin Perinato
encephaly. No
cranial defect near the foramen magnum and taken so that CSF appears d
Spinal ganglion

C D CHAPTER 17 |
F I G U R E 1 7 – 1 2 Diagrammatic sketches illustrate various types of spina bifida and the associated defects of the vertebral arches
(one or more), spinal cord, and meninges. A, Spina bifida occulta. Observe the unfused neural arch. B, Spina bifida with Neural tube
meningocele.
C, Spina bifida with meningomyelocele. D, Spina bifida with myeloschisis. The defects illustrated in B to D are referred to collectively
CHAPTER 17 | NERVOUS

as spina bifida cystica because of the cyst-like sac or cyst associated with them. CSF , Cerebrospinal fluid.
Neural tube
Neural fold

Neural fold

Rostral
neuropore
Rostral
neuropore Caudal

Somite
Somite

Defective closure of Defective closure of


Defective rostral
closureneuropore
of caudal neuropore
Defect
rostral neuropore caudal
1. Incomplete development Neural groove
F I G U R E 1 7 – 1 3 A fetus at 20 weeks with severe neural tube defects,
of brain with degeneration
Neural fold
1. 2. Incomplete
Incomplete development
development Neural groove
including acrania, cerebral regression (meroencephaly), iniencephaly (enlarge-
of calvaria
of brain 3.with degeneration
Alteration in facies (facial appearance)
ment of foramen
2. magnum), and a sacral dimple (arrow).
Incomplete +/–development
auricle
of calvaria
3. Alteration in facies (facial appearance)
Mass of brain
+/–tissue
auricle

Mass of brain
tissue
Neural deficit Unfused vertebral
caudal to lesion Meningomyelocele arch

Meroencephaly
+/– Clubfoot +/– Hydrocephalus Spina bifida occulta
Neural deficit Unfused ve
caudal to lesion Meningomyelocele Tuft of hair arch
Dura m
Skin

Meroencephaly Su
+/– Clubfoot +/– Hydrocephalus Spina bifida
Inc
ve
50 Tuft of
Myeloschisis Sp

Skin
Ve

Clubfoot

F I G U R EMyeloschisis
1 7 – 1 7 Schematic illustration shows the embryologic basis of neural tube defects. Meroencephaly
brain) results from defective closure of the rostral neuropore, and meningomyelocele results from defective clo
Defectos de nacimiento en el desarrollo del cerebro:
Microencefalia
• Trastorno del neurodesarrollo
• Calvaria y cerebro anormalmente pequeños
• Cara tamaño normal
• Deficiencia mental GRAVE à cerebro
subdesarrollado
• Origen: reducción en el crecimiento cerebral
• 25,000 bebés diagnosticados anualmente en
USA.
• Origen genético en algunos casos
• Exposición a radiación ionizante, agentes
infecciosos, alcohol y ciertos medicamentos
durante el período fetal pueden contribuir
• Diagnostico: US
Defectos de nacimiento en el desarrollo del cerebro:
Hidrocefalia
• Aumento significativo en el tamaño de la cabeza
• Causa: desequilibrio entre la producción y
absorción de líquido cefalorraquídeo (LCR)
• Consecuencia: exceso de LCR en el sist. ventricular
del cerebro
• El bloqueo de la circulación del LCR produce dilatación
de los ventrículos proximales a la obstrucción
• Esto ejerce cierta presión sobre los hemisferios
cerebrales (comprime el cerebro) à produce una tasa
acelerada de expansión del cerebro y el neurocráneo
• Síntomas
• Adelgazamiento de huesos de la calvaria
• Prominencia de la frente
• Atrofia de la corteza cerebral y la sustancia blanca
• Compresión de los ganglios basales y el diencpefalo
Defectos de nacimiento en el desarrollo del cerebro:
Holoprosencefalia
• Causa: separación incompleta de los hemisferios
cerebrales
• 1 de cada 250 fetos y 1 de cada 15,000 nacidos

• Factores genéticos y ambientales asociados


• Diabetes materna y teratógenos pueden destruir
las células embrionarias en el plano medio del
disco embrionario (3era semana) à amplia gama
de defectos congénitos que implican la formación
defectuosa del cerebro anterior
• Síntomas:
• Anomalías faciales
• Ojos anormalmente juntos
Defectos de nacimiento en el desarrollo del cerebro:
Hidrancefalia
• Anomalía poco común
• Ausencia de hemisferios cerebrales o
representados por sacos membranosos con
restos de corteza cerebral
• Causa: incierta, pero evidencia indica
obstrucción temprana de flujo sanguíneo
• Bebés parecen normales al nacer, pero la
cabeza crece excesivamente después del
nacimiento debido a la acumulación de LCR
• No ocurre desarrollo mental
• Poco o ningún desarrollo cognitivo
Defectos de nacimiento en el desarrollo del cerebro:
Malformación de Chiari
• Defecto estructural del cerebelo

• Proyección similar a la lengas de la médula y


desplazamiento inferior de la amígdala
cerebral a través del agujero occipital hacia el
canal vertebral
• Puede conducir a una hidrocefalia NO
comunicante
• Sistema ventricular distendido
Defectos de nacimiento en el desarrollo del cerebro:
Deficiencia mental
• Deterioro de la inteligencia
• Puede ser resultado de:
• Afecciones genéticas determinadas
• Gen mutante
• Anomalía cromosómica
• Abuso materno de alchool
• Grandes dosis de radiación
• Trastornos metabólicos
• Infecciones maternas (sífilis, rubéola, toxoplasmosis, citomegalovirus)
Referencias
1. Moore, K., Persaud, T., Torchia, M.(2016). The Developing Human:
Clinically Oriented Embryology. Elsevier. 10th edition. Philadelphia, PA.
2. Sadler, T. (2019). Langman´s Medical Embryology. Wlters Kluwer. 14th
edition. Philadelphia, PA.
3. Elshazzly, M. and Caban, O. (2018). Embryology, Central Nervous System.
Retrived from: https://www.ncbi.nlm.nih.gov/books/NBK526024/
4. Hill, M. (2019). Neural System Development. Retrived from:
https://embryology.med.unsw.edu.au/embryology/index.php/Neural_Syst
em_Development

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