EMBRIOLOGÍA ANIMAL

COMPARADA
PAOLA ESCOBAR RAMOS
BIÓLOGO
MSC. CIENCIAS EN BIOLOGIA
kar24239_ch05_161-211.qxd

HISTORIA DE VIDA
12/21/10 7:36 PM Page 162

Cleavage

Animal pole
n
io
at
liz Blastocoel
rti
Fe

G
as
tr
ul
at
io
n
Vegetal pole Blastula
Sperm
(male gamete)

Egg
(female gamete) Ectoderm
Gastrula Mesoderm
Endoderm
Adulthood

Gonad
Po velo and osi
de rva orph ies)
ste pm
(la tam pec
me st s

sis
mo

mb en

ne
ryo t

ge
nic

no
ga
Or
si
n

Ha
tch
ing

Histoge
nesis
 HISTORIA DE VIDA
kar24239_ch05_161-211.qxd 12/21/10 7:36 PM Page 162

• Desarrollo Cleavage

Animal pole

embrionario u
n
t io
l iza Blastocoel
rti
Fe

G
as
tr
ul
at
io
ontogenia

n
Vegetal pole Blastula
Sperm
(male gamete)

Egg

– Células en división
(female gamete) Ectoderm
Gastrula Mesoderm
Endoderm

Adulthood
• Maduración Gonad

1. Nacimiento
Po velo and hosi
de va orp ies)
ste pm
(la tam pec
me st s

sis
r
2. Madurez sexual
mo

mb en

ne
ryo t

ge
nic

no
ga
Or
si
n

Ha
tch
ing

Histoge
nesis

FIGURE 5.1 From single cell to millions of cells—life cycle of a frog. A sperm fertilizes the single-celled egg, and cell
division (cleavage) begins, leading to a multicellular blastula with a fluid-filled core (blastocoel). Major rearrangements (gastrulation) of
formative cellular layers (ectoderm, mesoderm, endoderm) lead next to an embryonic stage wherein these formative embryonic cells
become arranged into organs (organogenesis) and specific tissues (histogenesis). Upon hatching, the larva feeds and grows further,
eventually undergoing a major anatomical change (metamorphosis), becoming a juvenile and then an adult frog, which reproduces to
repeat the cycle.
 HISTORIA DE VIDA
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• Maduración Cleavage

Animal pole
n
io

1. Crecimiento
t
l iza Blastocoel
rti
Fe

G
as
tr
ul
at
io
n
2. Aprendizaje
Vegetal pole Blastula
Sperm
(male gamete)

Egg
(female gamete) Ectoderm

3. Carácteres físicos
Gastrula Mesoderm
Endoderm

Adulthood
distintivos Gonad

4. Juveniles
Po velo and hosi
5. Metamorfosis de va orp ies)
ste pm
(la tam pec
me st s

sis
r
mo

mb en

ne
ryo t

ge
nic

no
ga
Or
si

• Envejecimiento
n

Ha
tch
ing

Histoge
nesis

FIGURE 5.1 From single cell to millions of cells—life cycle of a frog. A sperm fertilizes the single-celled egg, and cell
division (cleavage) begins, leading to a multicellular blastula with a fluid-filled core (blastocoel). Major rearrangements (gastrulation) of
formative cellular layers (ectoderm, mesoderm, endoderm) lead next to an embryonic stage wherein these formative embryonic cells
become arranged into organs (organogenesis) and specific tissues (histogenesis). Upon hatching, the larva feeds and grows further,
eventually undergoing a major anatomical change (metamorphosis), becoming a juvenile and then an adult frog, which reproduces to
repeat the cycle.
 EMBRIOLOGÍA
• Aportes a la biologia evolutiva y
morfología
• Capas germinativas
– Endodermo
– Mesodermo
– Ectodermo
• Especies similares en ontogenia, son
evidencia de homologías lo que
indicaría una relación filogenética
 EMBRIOLOGÍA
• Zigoto
• Morula
• Blástula, blastoporo
• Gástrula,
arquenterón
• Néurula, sistema
nervioso
 EMBRIOLOGÍA

• Diferenciación del área embrionaria de la

extraembrionaria
• Capas germinativas, órganos
 EMBRIOLOGÍA
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• Historia de vida Cleavage

1. La fecundación
Animal pole
n
t io
l iza Blastocoel
rti
Fe

G
as
tr
ul
2. Desarrollo

at
io
n
Vegetal pole Blastula

embrionario
Sperm
(male gamete)

Egg
(female gamete) Ectoderm
Gastrula Mesoderm

3. Maduración
Endoderm

Adulthood
• Desarrollo Gonad

embrionario u
ontogenia Po velo and hosi
de va orp ies)
ste pm
(la tam pec
me st s

sis
r
mo

mb en

ne
ryo t

ge
nic

no
ga
1. Fecundación

Or
si
n

Ha
tch

2. Nacimiento o
ing

Histoge
nesis

eclosión FIGURE 5.1 From single cell to millions of cells—life cycle of a frog. A sperm fertilizes the single-celled egg, and cell
division (cleavage) begins, leading to a multicellular blastula with a fluid-filled core (blastocoel). Major rearrangements (gastrulation) of
formative cellular layers (ectoderm, mesoderm, endoderm) lead next to an embryonic stage wherein these formative embryonic cells
become arranged into organs (organogenesis) and specific tissues (histogenesis). Upon hatching, the larva feeds and grows further,
eventually undergoing a major anatomical change (metamorphosis), becoming a juvenile and then an adult frog, which reproduces to
repeat the cycle.
 EMBRIOLOGÍA
1. Fecundación n
tio
• Unión de gametos e r til
iz a
F
haploides
– Activa el
desarrollo Ve
embrionario Sperm
(male gamete)
• El huevo acumula Egg
vitelogenina (female gamete)

– Vitelo
thood
 EMBRIOLOGÍA:
Fecundación
• Unión de los gametos maduros
– Espematozoide y óvulo
– Maduros
– Haploides
• Óvulo acumula una gran cantidad de
vitelo, nutrientes que soportaran al
embrión
• Cantidad de vitelo específico para cada
especie
radiata is not a separate layer but an effect produced by a
EMBRIOLOGÍA:
dense stand of microvilli projecting from the surface of the
ovum. These microvilli intermingle with microvilli reaching
Tipos De Huevos De Acuerdo A La
Cantidad De Vitelo
TABLE 5.1 Comparison of Cleavage Patterns
and Yolk Accumulation in Representative Vertebrates
Cleavage Yolk Representative
Pattern Accumulation Animals
Holoblastic Microlecithal Amphioxus, eutherian mammals
Mesolecithal Lampreys, bowfins, gars, amphibians
Meroblastic Macrolecithal Elasmobranchs, teleost fishes
Discoidala Macrolecithal Reptiles, birds, monotremes

a
Discoidal cleavage is an extreme case of meroblastic cleavage.
 EMBRIOLOGÍA
• De acuerdo a la
distribución del vitelo
– Isolecitos, distribuídos
uniformemente
– Telolecitos, distribuídos
en un polo
– Centrolecítos,
distribuídos en el centro
 EMBRIOLOGÍA
• De acuerdo al Cleavage

arreglo irregular
del vitelo,n Animal pole
tio
polaridad
ili
z a
Bl
t
er
– Polo
F vegetal, mayor
cantidad de vitelo
– Núcleo haploide
prominente, polo Vegetal pole Blastula
animal
erm
ale gamete)
 EMBRIOLOGÍA:
Estructura Del Huevo

• Cortex, contiene granulos corticales

• Membrana plasmática
• Tres membranas:
 EMBRIOLOGÍA:
Estructura Del Huevo

• Membrana primaria del huevo, membrana

vitelina o Zona pelucida
 EMBRIOLOGÍA:
Estructura Del Huevo

• Membrana secundaria del huevo, células

foliculares (corona radiante)
– Rodean al huevo
– Transfieren nutrientes
 EMBRIOLOGÍA:
Estructura Del Huevo

• Membrana terciaria del huevo, en el

oviducto
• Envoltura tiburones
• Cáscara aves, reptiles y monotrema
 EMBRIOLOGÍA

• Oviparos, huevo al nacer

• Incubación
• Viviparos, vivo al nacer
• Segmentación, posterior a la fecundación
(b) Amphibians

EMBRIOLOGÍA: Segmentación
kar24239_ch05_161-211.qxd 12/21/10 7:36 PM Page 165
Blastoderm
Subgerminal space

(c) Reptiles/birds

• Clivaje Zygote
Inner cell mass
Morula Blastula Sagittal section of blastula

– Morula FIGURE 5.2 Cleavage stages

in five chordate groups. Relative
sizes are not to scale. (a) Amphioxus.
– Blastula
(d) Mammals
(eutherian)
(a) Amphioxus (b) Amphibian. (c) Reptiles and birds.
(d) Eutherian mammal. Blastoderm
• Blastómeros
• Holobástico,
wedges. The third cleavage plane is at right angles to the first
two and lies between poles just above the equator, produc-
huevos
ing the eight-celled morula stage (figure 5.2a). Subsequent
divisions of the blastomeres, now less and less in synchrony
(b) Amphibians

microleciticos
with each other, yield the 32-celled blastula surrounding the
fluid-filled blastocoel.
Blastoderm
Subgerminal space

Fishes
– Anfioxos (c) Reptiles/birds

In gars and– Mamíferos

bowfins, euterios
cleavage is holoblastic, although cleav-
FIGURE 5.3
Amia.
Holoblastic cleavage in theInner
bowfin,
cell mass

age furrows of the vegetal pole are slowed. Most cell division
After Korschelt.
is restricted to the animal pole (figure 5.3). Blastomeres in
the vegetal pole are relatively large and hold most of the yolk
reserves; those in the animal pole are relatively small and Amphibians
(d) Mammals
form the blastoderm, a cap of cells arched over a small blas-
(b) Amphibians

EMBRIOLOGÍA: Segmentación
kar24239_ch05_161-211.qxd 12/21/10 7:36 PM Page 165
Blastoderm
Subgerminal space

(c) Reptiles/birds

• Holobástico, surcos
Inner cell mass
Zygote Morula Blastula Sagittal section of blastula

mitóticos FIGURE 5.2 Cleavage stages

in five chordate groups. Relative
sizes are not to scale. (a) Amphioxus.
perpendiculares
(d) Mammals
(eutherian)
(a) Amphioxus (b) Amphibian. (c) Reptiles and birds.
(d) Eutherian mammal. Blastoderm

• Bola hueca llena de

wedges. The third cleavage plane is at right angles to the first
líquido, blastocele
two and lies between poles just above the equator, produc-
ing the eight-celled morula stage (figure 5.2a). Subsequent
divisions of the blastomeres, now less and less in synchrony (b) Amphibians
with each other, yield the 32-celled blastula surrounding the Blastoderm
fluid-filled blastocoel. Subgerminal space

(c) Reptiles/birds
Fishes
FIGURE 5.3 Holoblastic cleavage in the bowfin,
Inner cell mass
In gars and bowfins, cleavage is holoblastic, although cleav-
Amia.
age furrows of the vegetal pole are slowed. Most cell division
After Korschelt.
is restricted to the animal pole (figure 5.3). Blastomeres in
the vegetal pole are relatively large and hold most of the yolk
reserves; those in the animal pole are relatively small and Amphibians
(d) Mammals
form the blastoderm, a cap of cells arched over a small blas-
 EMBRIOLOGÍA: Segmentación
• Huevos
macroleciticos,
división kar24239_ch05_161-211.qxd 12/21/10 7:36 PM Page 166

meroblastica
– Solo una porción Blastoderm
Blastocoel

del citoplasma se Vitelline

membrane
Periblast

segmenta (a) (b) (c) (d)

Yolk mass

FIGURE 5.4 Discoidal cleavage in a teleost (zebra

fish). (a) Cleavage begins with the appearance of the first mitotic
furrow.After successive mitotic divisions (b), the blastula (c) results.
(d) Cross section of the blastula. A cap of blastoderm rests on the
uncleaved yolk mass, and a vitelline membrane is still present
around the entire blastula.
After Beams and Kessel in Gilbert.

(future embryo) and a hypoblast (supportive structures).

Yolk is so prevalent within the vegetal pole that cleavage
 EMBRIOLOGÍA: Segmentación
• Segmentación
meroblástica
discoidal kar24239_ch05_161-211.qxd 12/21/10 7:36 PM Page 166

– Segmentación
restringida al polo Blastoderm

animal
Blastocoel
Vitelline
membrane
Periblast

– Aves, reptiles, (a) (b) (c) (d)

Yolk mass

peces y FIGURE 5.4 Discoidal cleavage in a teleost (zebra

fish). (a) Cleavage begins with the appearance of the first mitotic
furrow.After successive mitotic divisions (b), the blastula (c) results.

monotremas (d) Cross section of the blastula. A cap of blastoderm rests on the
uncleaved yolk mass, and a vitelline membrane is still present
around the entire blastula.
After Beams and Kessel in Gilbert.

(future embryo) and a hypoblast (supportive structures).

Yolk is so prevalent within the vegetal pole that cleavage
 EMBRIOLOGÍA: Segementación
en el Anfioxo
Zygote Morula Blastula Sagittal section of blastula

(a) Amphioxus

Blastoderm
• Huevo microlecítico
• Surcos mitóticos del polo animal al vegetal
• Mórula de 4 a 8 células
• Blástula
(b) Amphibians de 32 células
Blastoderm
Subgerminal space
 FIGURE 5.2 Cleavage stages

EMBRIOLOGÍA: Segementación in five chordate groups. Relative

sizes are not to scale. (a) Amphioxus.
(b) Amphibian. (c) Reptiles and birds.

en Peces (d) Eutherian mammal.

o the first
r, produc-
ubsequent
ynchrony
nding the

FIGURE 5.3 Holoblastic cleavage in the bowfin,

ugh cleav-
Amia.
ll division
After Korschelt.
omeres in
• Segmentación
of the yolk
small and Amphibians
holoblástica
mall blas-
• Mayor
e that of
divide división restringida
those of the vegetalal polo
As in the gars and bowfins, blastomeres of the animal pole
more often than pole, animal
in which
cell division is presumably slowed by abundant yolk
teleosts,
•
olky cyto-
eosts pro-
Polo vegetal con blastómeros
are larger than those ofmas
platelets. Consequently, cells of the vegetal pole, having
undergone fewer divisions, the more
hese is the
a discrete
grandes y conof the
small blastomeres mayor
animal polevitelo
active animal pole. When the blastula stage is reached, the
constitute the blasto-
derm and form a roof over the emerging blastocoel.
cause it is
a–d). The
ytial layer Reptiles and Birds
EMBRIOLOGÍA: Segmentación en
(a)
Anfibios
Amphioxus

Blastoderm

(b) Amphibians
Blastoderm

• Similar a los peces Subgerminal space

• Blastomeros conforman el blastodermo y

(c) Reptiles/birds

Inner cell mass

una cubierta sobre el bastocele
FIGURE
in five chor
sizes are not
(d) Mammals (b) Amphibia
(eutherian) (d) Eutherian

wedges. The third cleavage plane is at right angles to the first

 EMBRIOLOGÍA: Segementación
en Peces
Blastoderm yolk. T
Blastocoel yolk an
Vitelline
membrane
Marsup
Periblast
modest
Yolk mass
(a) (b) (c) (d) by a zo
corona
FIGURE 5.4 Discoidal cleavage in a teleost (zebra the ovi
fish). (a) Cleavage begins with the appearance of the first mitotic shell m
• Mixinis y condrictios, segmentacion
furrow.After successive mitotic divisions (b), the blastula (c) results.
(d) Cross section of the blastula. A cap of blastoderm rests on the
it is ge
compos
discoidal
uncleaved yolk mass, and a vitelline membrane is still present
around the entire blastula.
and so
secreted
• Dos tipos de células:
After Beams and Kessel in Gilbert. and ute
cleavag
Blastodermo:
– (future cuerpo del embrión
embryo) and a hypoblast (supportive structures).
support
tion, it
Periblasto:
– Yolk moviliza
is so prevalent elvegetal
within the vitelo pole that cleavage
Ea
furrows do not pass through it at all; thus, cleavage is dis-
tion of
coidal. Blastomeres resulting from successful cleavage
 EMBRIOLOGÍA
(b) Amphibians
Blastoderm
Subgerminal space

(c) Reptiles/birds

Inner cell mass

• Polo vegetal abundante, surcos no lo FIGURE 5.2

atraviesan in five chordate
sizes are not to sc
(d) Mammals (b) Amphibian. (c)
• Blastómeros en el polo animal y forman
(eutherian) (d) Eutherian mam

delwedges.
blastodermo
The third cleavage plane is at right angles to the first
two and lies between poles just above the equator, produc-
• Espacio subgerminal,
ing the eight-celled espacio lleno de
morula stage (figure 5.2a). Subsequent
divisions of the blastomeres, now less and less in synchrony
liquido entre
with each other, el blastodermo
yield the 32-celled
fluid-filled blastocoel.
blastula surrounding the y vitelo
Fishes
EMBRIOLOGÍA: Segmentación en
Mamíferos Monotremas
Zona pellucida Morula Blastocyst
Blastocoel
Blastoderm

Yolk mass

(a) Monotreme

• Blástula en
Yolkmamiferos:
mass Blastocisto FIG
Zona pellucida Blastocoel
• Blastodermo crece hasta envolverZonaelpellucida livin
cleav
Mucoid layer
vitelo Protoderm
blast
cleav
(b) Marsupial Shell membrane comp
prod
 Blastocoel
Blastoderm

EMBRIOLOGÍA: Segmentación en
Yolk mass
Mamíferos Marsupiales
(a) Monotreme

Yolk mass
Zona pellucida Blastocoel FIGURE
Zona pellucida living mam
Mucoid layer cleavage, wit
blastoderm a
Protoderm
cleavage doe
(b) Marsupial Shell membrane composed o
produced du
the zona pel
• Sin morula Zona pellucida protoderm.T
Inner cell mass thin shell me
• Blastomeros rodea a la zona pelúcida Trophoblast y
from a moru
aside as an in
forman un protodermo Blastocoel
trophoblast)
coat and zon
Follicle cells
morula but a
• Un (c)polo forma el embrión
(corona radiata)
Eutherian mammal Monotreme base

terms choriovitelline membrane and chorioallantoic membrane just one part of the em
 (b) Amphibians

EMBRIOLOGÍA: Segmentación en Blastoderm

Subgerminal space

Mamíferos Euterios
(c) Reptiles/birds

Inner cell mass

FIG
in fiv
sizes
(d) Mammals (b) Am
(eutherian) (d) Eu

• Morula
wedges. The third cleavagebola
plane is atde blastomeros
right angles to the first
two and lies between poles just above the equator, produc-
• Formación
ing the eight-celled del
morula blastocele y luego el
stage (figure 5.2a). Subsequent
blastocisto
divisions of the blastomeres, now less and less in synchrony
with each other, yield the 32-celled blastula surrounding the
• Blastocisto
fluid-filled blastocoel. membranas adicionales
yFishes
extraembrionarias y cuerpo del embrión
FIGURE 5.3 Holoblas
In gars and bowfins, cleavage is holoblastic, although cleav-
Amia.
age furrows of the vegetal pole are slowed. Most cell division
 EMBRIOLOGÍA:
Gastrula y Neurula
• Gastrula, tubo digestivo primitivo
– Gastrocele o arquenteron
• Neurula, tubo neural
kar24239_ch05_161-211.qxd 12/21/10 7:36 PM Page 168

• Gastrula y neurula ocurren

simultáneamente
• Capas germinativas ocupan sus Notochord
Neural plate

posiciones iniciales Ectoderm

Mesoderm

(a)
Endoderm Gastrocoel
 kar24239_ch05_161-211.qxd 12/21/10 7:36 PM Page 170

EMBRIOLOGÍA:
Gastrula y Neurula en Anfioxo
• Gastrula,
invaginación de la Blastocoel

pared vegetal P

• Neurula, se P

P
delimitan (a) Gastrocoel

mesodermo y Blastopore

ectodermo
 EMBRIOLOGÍA:
kar24239_ch05_161-211.qxd 12/21/10 7:36 PM Page 170

Gastrula y Neurula en Anfioxo

Chordamesoderm

Neural plate
Blastocoel
Early
Gastrocoel
P

• Mesodermo se P(a)
P

evagina, celoma
Gastrocoel

Blastopore

Ectoderm

• Mesodermo se Mesoderm

diferencia en Endoderm

cordomesodermo,
notocorda FIGURE 5.7 Amphioxus gastrulation and
neurulation. (a) At the left, invagination at the vegetal pole Neural tube
pushes cells into the interior of the blastula.The blastocoel is Somite
eventually obliterated, and the new space these ingrowing cells Coelom
define becomes the gastrocoel.The arrow indicates the
anteroposterior axis of the embryo. (b) Successively older cross Notochord
sections taken along the plane (P) defined in the illustration on
the left (a). As development proceeds, mesodermal outpocketings Gut Late
appear and pinch off to form somites, leaving the endoderm to
form the lining of the gut. (b)