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Primeras ballenas
• Endoesqueleto, vértebras
• Crecimiento
• Faringe perforada
• Apéndices pareados
CORDADOS: CARACTERÍSTICAS QUE LOS
DISTINGUEN
• Aparición, período Cámbrico 530 millones de años
PHYLUM CHORDATA: Notochord
• Primeros cordados, cuerpo suave
In a lancelet, the simplest chordate, the
flexible notochord persists throughout
life and aids swimming by giving muscles
Notochord
1. Notocordio
Water
Lancelets lack pigment in their
Dorsal nerve skins, and so are transparent.
cord
Oral hood
with tentacles
3. Hendiduras faringeas
sensory systems. The
Atriopore
animal has no head, eyes,
ears, or nose. Instead,
sensory cells that detect
chemicals line the oral Anus
Intestine
tentacles.
The Nonvertebrate Chordates The tadpolelike larvae of tunicates plainly exhibit all
the basic characteristics of chordates and mark the tun
• Endoesqueleto; locomocion, soporte y protección
Tunicates
The tunicates (subphylum Urochordata) are a group of
cates as having the most primitive combination of featur
found in any chordate (figure 48.5c). The larvae do n
mals. In colonial tunicates, there may feed and have a poorly developed gut. They remain fre
about 1250 species of marine animals. Most of them are
be a common sac and a common swimming for only a few days before settling to the bo
sessile as adults (figure 48.5a,b), with only the larvae hav-
opening to the outside. There is a tom and attaching themselves to a suitable substrate b
ing a notochord and nerve cord. As adults, they exhibit
Marinos
gonads, indicating that these are maleGenital
lancelets. Tunic
and persists throughout the animal’s
•
duct
life.
Lancelets spend most of their time Gonad
Stomach
partly buried in sandy or muddy substrates, with Oral hood Notochord Dorsal Heart
with tentacles
only their anterior ends protruding (figure 48.6). nerve cord Intestine
They can swim, although they rarely do so. Their (b)
Branchial
pouch Proboscis
Branchial coelom
kar24239_ch02_048-081.qxd 12/17/10 6:43 PM Page pore
60
Mouth Stomochord
Trunk Collar
coelom coelom
HEMICHORDATA
FIGURE 2.8 Hemichordate, generalized acorn worm. Proboscis, collar, and trunk regions are shown in partial cutaway view,
revealing the coelom in each region and the associated internal anatomy of the worm. Within the proboscis is the stomochord, an
Tentacles
extension of the digestive tract. The food-laden cord of mucus
Collar(spiral arrow at right) enters the mouth together with water. Food is
directed through the pharynx into the gut. Excess water exits via the pharyngeal slits. Several slits open into each branchial pouch, a
Proboscis
common compartment with a branchial pore that opens to the outside environment. Arms
Blastocoel in length, but most are shorter than this. Most live in mucus-
lined burrows and have a body with three regions—proboscis,
Coelomic
Nephridium compartment
Oral collar, trunk—each with its own coelom (figure 2.10a–c).
lamella
Nephridial ductArms
Mouth The proboscis,
Tube
used in both locomotion and feeding, includes
Preoral a muscular outer wall that encloses a fluid-filled coelomic
hood space. Muscular control over the shape of the proboscis gives
Zooid
Stalk
the animal a useful probe to shape a tunnel or inflate itself
Gut
Trunk against the walls of the burrow to anchor its body in place
Nephridial pore (figure 2.10b). Tucked away in their burrows, many species
Pulsatile vesicle
Circumoral ingest loosened sediment, extract the organic material it con-
band
tains, pass the spent sediment through their simple gut, and
Telotroch deposit
(a) (b) a casting (fecal waste) on the surface of the substrate
where changing tides flush it away. Some wide-bodied,
FIGUREFIGURE 2.9 Hemichordate,
2.14 Hemichordata,generalized tornaria (a) The sessiledeep-sea
Pterobranchia. pterobranchenteropneusts crawl
Rhabdopleura. Notice thatand glide along
this pterobranch the abyssal
has the
larva. The same
simple gutplan
body begins
as anatacorn
the mouth under a preoral
worm—proboscis, hood these three
collar, trunk—but oceanfeatures are modified and the whole animal lives in a tube.
bottom.
(a)
Dorsal and tubular
Myomeres nerve cord
Dorsal fin Ocellus
Notochord
Caudal fin
Hindgut Atriopore
Anus Midgut cecum
Ileocolic ring
Esophagus Oral hood
Postanal tail Midgut Pharyngeal slit
(b)
CEPHALOCHORDATA
Ocellus Notochord Wheel organ
Dorsal and Velar tentacle
tubular nerve cord Velum
Muscles Right
Testes stomach
lobe
Incurrent Ovary Ovary
Cecum
Oral Pharynx Testes
siphon (mouth) Intestine Intestine Left
tentacle
stomach
Dorsal lamina Cilated funnel Mouth Trunk
lobe
Right
Endostyle
Ganglion Anus stomach Esophagus
Incurrent lobe
Heart Pharyngeal
Subneural gland Anus slit
Pharyngeal
slit Prebuccal
Atrial Endostyle gland
Excurrent Excurrent
siphon opening Mouth
(atrial) Feeding-
Screen
siphon filter
Tunic Gelatinous
matrix
Atrium (“House”)
Tail
(a)
Stigmata
Dorsal
(b)
Anus
(c)
UROCHORDATA
Heart FIGURE 2.26 Urochordata, Larvacea (Appendicularia), Oikopleura albicans. (a) Oikopleura is shown within its gelat
house.The animal’s feeding-filter obtains food from the incurrent of circulating water (small arrows).This larvacean resides at the ba
Gonad
Ciliated band the screen where it sucks food off these screens. (b and c) Enlarged lateral and ventral views, respectively, of the isolated larvacean. T
undulating tail, supported by a notochord, is active in producing the current of food-bearing water that moves through the internal
Stomach
channels of the house and through the feeding-filter.
(a) After Flood; (b, c) after Alldredge.
Stigmata
(b) Vessels of
branchial
new house continues with the addition of feeding screens feeders. These tiny organisms that escape the clutc
basket
and filters. Sometimes within the space of only a few min- other suspension feeders fall prey to the efficient fil
Dorsal lamina utes, the new house is complete and the larvacean is once gear of larvaceans. Larvacea are able to sift through
again feeding actively. volumes of water, ingest a wide range of plankton
All species, except one, are monoecious, and most of including the very tiniest, and proliferate rapidly in res
these are protandrous; that is, sperm and eggs are produced to local blooms in food supply.
Organs
by the same gonad (of the same individual) but at different The trunk of the larvacean holds its major body o
Heart times during its life. Maturation is so rapid that within 24 to although which organs are present varies among the
Posterior 48 hours of fertilization, miniature larvaceans secrete a
Subendostyle families of Larvacea. Members of the smallest famil
vessel vessel house and are set up for feeding. Kowalevskiidae, lack endostyle and heart. In the Fritilla
(d) Their rapid reproduction and special feeding appara- the stomach consists of only a few cells. In the Oikop
CORDADOS: CARACTERÍSTICAS QUE LOS Kardong: Vertebrates: 2. Origin of Chordates Text
• Mesodermo br
Connective tissue m
sheath
ve
• Encima del celoma, debajo del sistema Collagenous on
sheath it
nevioso central fo
Vacuole in fu
notochordal
cell
na
• Compuesta por: m
ic
• Invaginación
56 Kardong: Vertebrates: 2. Origin of Chordates Text © The McGraw−Hill
Comparative Anatomy, Companies, 2009
Function, Evolution, Fifth
notocorda
and cilia, also becomes the basis for procurement and pro- tary canal and is a tube (although the
cessing of large food items. Slits still serve as convenient exit
FIGURE 2.6 Dorsal hollow nerve cord. (a) Basic body
plan of an annelid or arthropod. In such animals, a definitive nerve
cord, when present, is ventral in position, solid, and lies below the
portals for excess or spent water, while adjacent gill struc-
tures function in respiration. In fishes and aquatic amphib- hollow center may be nearly obliter-
digestive tract. (b) Basic chordate body plan.The nerve cord of
chordates lies in a dorsal position above the digestive tract and
ians, the pharyngeal slits that appear during embryonic notochord. Its core is hollow, or more correctly, it has a fluid-
development usually persist into the adult and form the exit ated during growth). The anterior end
filled central canal, the neurocoel, indicated as the white spot in
• Neurocele channel through which water associated with feeding and the dorsal hollow nerve cord.
respiration flows. In vertebrates that reside on land, how-
ever, the embryonic pharyngeal slits normally never open
becomes enlarged to form the brain.
and thus do not give rise to any adult derivative. form the central nervous system do not amass as thickened
Why cilia are replaced by muscles
The hollow cordinstead,iscellsproduced
surface plates (placodes); individually move in the
inward to assemble into the basic nervous system. Further, the
as body size increases (p. 128)
embryo majorby the
nerve cord in mostinfolding ofposition,
invertebrates is ventral in
below the gut, and solid. In chordates, however, the nerve
ectodermal
CORDADOS: CARACTERÍSTICAS QUE LOS
DISTINGUEN - HENDIDURAS FARÍNGEAS
• Faringe parte del tubo digestivo
• Función respiración
• Alimentación
Incurrent
Oral
Endostyle siphon (mouth)
tentacle
Dorsal lamina Cilated funnel
Subneural gland
partículas
Atrial
and Slits
Excurrent
siphon
(atrial)
Ventral siphon
blood Tunic
sinus
Dorsal
Grupo de cilios
(a) Endostyle
• Stigmata
Gonad
Ciliated band
Stomach
Stigmata
(b) Vessels of
branchial
basket
Dorsal lamina
(c) Organs
Heart
Posterior Subendostyle
tebrates, the nerve
tostomes. Embryological cord passes
characteristics, through
modern molecular
phylogenies, and the fossil record all imply that there was
rigid inte
skin. Th
theanprotective neural
ancient and fundamental arches
divergence betweenof
the the
proto- ver-
port, and
Gut Notochord
Postanal tail
Atrium
Pharyngeal slit in pharynx
Pharyngeal slits
(c)
are perforated
(b)
slitlike
openings that Dorsal
leadhollow from the pharyn-
• Metabolismo del yodo geal cavity to the
nerve cord*
outside. They are
Notochord*
formed by the Gut
inpocketing of the out- FIGURE 2
side ectoderm (pharyngeal
Pharynx* grooves)
(a) A single str
pharynx, and t
Atrium
lower chordat
Coelom common enclo
the single atrio
Endostyle*
the floor of th
Body wall showing the tu
(c) Cross sect
(c) (b) indicate chord
(sharks)
CORDADOS: CARACTERÍSTICAS QUE LOS
⎯⎯⎯⎯ Chordata ⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯→
DISTINGUEN - RABO O COLA POSTANAL
⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯ Vertebrata ⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯→
⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯ Craniata ⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯→
⏐ ←⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯ Gnathostomata ⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯→
⏐ ←⎯⎯⎯⎯ Pisces ⎯⎯→ ⏐ ←⎯⎯⎯⎯⎯⎯⎯ Tetrapoda ⎯⎯⎯⎯⎯⎯⎯→
⎯⎯⎯ larvas de tunicados⏐ y←⎯⎯⎯ Amniota ⎯⎯⎯⎯⎯→
Motilidad,⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯→
• Anamniota
anfioxus
• Propulsión,
e evagination, peces
or outpocketing, Segmented myotomes between
endodermal lining of the phar- septa anchored to notochord
• Coxis,
haryngeal humanos
pouches). In aquatic
tes, the two pockets break
h the pharyngeal cavity where
• Rabo, mamíferos
eet to form the pharyngeal slit.
Postanal tail
iotes these pockets may not
• Prolongación
hrough the pharyngealdelcavity
cuerpo
ly grooves are formed instead Postanal Tail
In tetrapod (four-footed) verte- The postanal tail, together with so-
the pharyngeal pouches give matic musculature and the stiffening
HISTORIA EVOLUTIVA DE LOS CORDADOS
• Cefalización
Nerve ganglion
Pharynx
Organismo filtrador
Excurrent
• siphon
Endostyle
Gill slit
Intestine
• Forma de tunicado Genital duct
Tunic
Gonad
Stomach
Heart
Mouth Atriopore
(incurrent siphon) (excurrent siphon)
Pharynx
with gill slits Stomach
Meiofauna o filtradores
Notochord
• (c) Heart
FIGURE 48.5
Tunicates (phylum Chordata, subphylum Urochordata).
(a) The sea peach, Halocynthia auranthium. (b) Diagram of the
structure of an adult tunicate. (c) Diagram of the structure of a
larval tunicate, showing the characteristic tadpolelike form. Larval
tunicates resemble the postulated common ancestor of the
ar24239_ch02_048-081.qxd 12/17/10 6:44 PM Page 75
Kardong: Vertebrates: 2. Origin of Chordates Text © The McGraw−Hill 53
Cordados
The first person toprovenientes debody
anélidos
pore being the first). The now multicellular embryo is com-
• propose that the chordate plan was
Embryonic development; details of early cleavage (p. 161)
posed of three basic tissue layers: an outer ectoderm, an inner
In both bilaterian groups, the egg begins to divide endoderm that forms the lining of the gut, and a mesoderm
Gut after fertilization, a process termed cleavage, until
derived from a flipped-over version of an arthropod’s was repeatedly
Nerve
the very young embryo cord
is made up of many cells formed from
that forms the layer between the two. If a solid mass of meso-
Annelid/Arthropod
dermal cells splits to form the body cavity within them, the
Geoffroy Saint-Hillaire, French zoologist. In 1822 he put the original single-celled egg (figure 2.3). In some animals, result is a schizocoelom (figure 2.3a). If, instead, the mesoderm
•forthArgumentación débil
this theory, perhaps inspired by his dissections of lob-
dividing cells of the embryo are offset from each other, a pat-
tern known as spiral cleavage. In others, the dividing cells are
arises as outpocketings of the gut that pinch off to form the
body cavity, the result is an enterocoelom (figure 2.3b).
sters, but also as part of his larger view that all animals
shared a common, underlying body plan that nature Coelom Mesoderm
Comparaciones sobre
Georgesanalogías
Protostome
•molded into dramatic variations. Cuvier led those Blastopore Mouth
Annelid
(earthworm)
Anus
•gist El
W. cordón nervioso
H. Gaskell and, shortly thereafter, William Patten
(modern bony fish)
Primitive Anus
revived the issue and presented a closely argued case in sup- Deuterostome
gut
Coelom
• Desarrollo
Their reasoning,detaken
la boca
together, went as follows. Fish Echinoderm Mouth
present in annelids and arthropods, although upside down Mammal Anus develops
from blastopore
(figure 2.29a,b). In annelids and arthropods, the nerve cord FIGURE 2.3 Protostomes and deuterostomes. Bilateria are divided into two major groups on the basis of embryonic
occupies a ventral position below the gut along with a major characteristics. (a) Protostomes usually show spiral cleavage, coelom formation by splitting of the mesoderm, and derivation of the mouth
(b)
from the blastopore. (b) Deuterostomes often exhibit radial cleavage, coelom formation by outpocketing of the gut, and derivation of the
HISTORIA EVOLUTIVA DE LOS CORDADOS
• Cordados provenientes de los equinodermos
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Garstang reasoned that because of these embryonic affini- they met at the midline together with the underlying nerve
ties, echinoderms or a group very similar to echinoderms tract, the antecedent of the nerve tube (figure 2.30).
• Banda circumoral, locomoción
were the likely chordate ancestors.
At first this seemed farfetched. Adult echinoderms, such
Garstang pointed to the rolling up of the neural tube during
vertebrate embryology as an embryonic remnant of this
as starfish, sea urchins, sea cucumbers, and crinoids, offer little phylogenetic event. The segmental tail musculature with
to suggest a phylogenetic affinity with chordates. They have notochord evolved simultaneously with the neural tube—
tube feet, calcium carbonate plates in their skin, and pentara- essentially a new nervous system serving to activate and
dial (five-armed) body symmetry. Let’s examine Garstang’s control the new muscularized tail. On the other hand,
HISTORIA EVOLUTIVA DE LOS CORDADOS
kar24239_ch02_048-081.qxd 12/17/10 6:44 PM Page 76
Similitudes bioquímicas en las proteínas chordates (left) was bilaterally symmetrical Circumoral Dorsal nerve
• and had the external appearance of a ciliated band cord Pharyngeal
slits
young echinoderm larva. The ancestor’s
sanguíneas circumoral ciliated bands and their
associated underlying nerve tracts moved
Mouth
Ciliated Endostyle
dorsally to meet and fuse at the dorsal adoral band
midline, forming a dorsal nerve cord in the Gut
• Química muscular: compuestos de fósforo chordate body plan. The adoral ciliated band
gave rise to the endostyle and ciliated tracts
within the pharynx of the chordate.
Scientists other than Garstang noted that Notochord
• Equinodermos y pterobranquios
relacionados con su ancestro
• Larva placntónica
• Cuerpo bipartito:
• Esqueleto interno
HISTORIA EVOLUTIVA DE LOS CORDADOS
• Swalla y Smith (2008)
• Notocorda dorsal
• Cola postanal
• Argumento no consensuado
HISTORIA EVOLUTIVA DE LOS CORDADOS
• Yunnanozoons, cordado basal
• Semejante a un pez
• Cola postanal
• Notocorda
• Branquias
• Miómeros laterales
• Conodontos
• Alargados
• Cuerpo suave
• Paquetes musculares en V
• Ojos
• Denticulos conicos de
fosfato de calcio
HISTORIA EVOLUTIVA DE LOS
VERTEBRADOS
• Remodelaciones corporales
• Ventajas fisiológicas
• Adaptaciones favorables
evolución de los vertebrados.
HISTORIA EVOLUTIVA DE LOS VERTEBRADOS:
Paedogenesis puede ocurrir en la evolución de los
vertebrados
• Paedogenesis Adultos conserva características
Paedogenesis: Adultos conserva características
que se observan en el desarrollo embrionario
que se observan en el desarrollo embrionario
Vertebrate
No Paedogenesis Paedogenesis
Arthropods
Sponges
Flatworms
Annelids
Chordates
Nematodes
Cnidarians
Mollusks
Echinoderms
y aereos
the echinoderms, the only other
uterostomes. However, unlike
hinoderms, chordates are character-
Diferentes estilos de vida
d by a• notochord, jointed appendages,
d segmentation. There are some
000 species of chordates, a phylum
t includes birds, reptiles, amphib-
Características de los cordados
• and mammals.
s, fishes,
Four features characterize the chor-
es and have played an important
Al igual que los tunicados y
• evolution of the phylum
e in the
gure 48.2):
anfioxus Postanal
tail
1. A single, hollow nerve cord
Notochord
runs just beneath the dorsal sur-
face of the animal. In verte-
• Cinco características del filum
brates, the dorsal nerve cord differentiates into the
FIGURE 48.2
Some of the principal features of the
brain and spinal cord. chordates, as shown in a generalized
2. A flexible rod, the notochord, forms on the dorsal embryo.
side of the primitive gut in the early embryo and is
SUBFILUM VERTEBRATA: COLUMNA Y
CRÁNEO
48.3 The vertebrates have an interior framework of bone.
• La notocorda es reemplazada
Characteristics
durante el desarrollo of Vertebrates
Ectoderm
Vertebrates (subphylum Vertebrata) are chordates with a
embrionario
spinal column. The name vertebrate comes from the indi- Neural tube
vidual bony segments called vertebrae that make up the
Notochord
spine. Vertebrates differ from the tunicates and lancelets in
• Columna vertebral ósea
two important respects:
1. Vertebral column. In vertebrates, the notochord is
replaced during the course of embryonic develop-
• Segmentos que cubren y
ment by a bony vertebral column. The column is a
series of bones that encloses and protects the dorsal
Vertebral
body developing
Forming
protejen el nervio dorsal
nerve cord like a sleeve (figure 48.8).
2. Head. In all vertebrates but the earliest fishes, there
around notochord
neural arch
Blood vessels
is a distinct and well-differentiated head, with a skull
and brain. For this reason, the vertebrates are some-
• Cabeza diferenciada: esqueleto
times called the craniate chordates (Greek kranion,
“skull”).
y Incerebro
addition to these two key characteristics, vertebrates
differ from other chordates in other important respects: Rib Neural arch
Notochord
spine. Vertebrates differ from the tunicates and lancelets in
two important respects:
• Derivan del tubo
1. Vertebral column. neural
In vertebrates, the notochord is
replaced during the course of embryonic develop-
ment by a bony vertebral column. The column is a Vertebral
• Espina dorsal
• Notocordo
• Arco neural
• Médula espinal
Neural spine Dorsal hollow
nerve cord
• Arco hemal Intervertebral Neural arch
body
Centrum
• Vasos sanguíneos
Hemal arch Blood vessel
• Primeros vertebrados, sin vertebras
Hemal spine
pero con notocordo fuerte
Mesoderm
(a)
Epimere
crestas neurales, caracteres Mesomere
• No observados en los
ancestros
Neural crest Neurocoel
Neural tube
Chordamesoderm
Organos sensoriales y
Dermatome
• Somite Myotome
Neural plate
algunos dientes
Dorsal mesentery
Blastocoel Sclerotome
Early
GastrocoelSomatopleure
(c) Gut tube cavity
P Splanchnopleure
• Epidermis
• Epitelio estratificado
• Dermis
• Tejido conectivo
SUBFILUM VERTEBRATA: Característica diagnósticas
de los cordados en alguna etapa de su vida
• Faringe musculosa
• Cilios
• Agnathos
• Gnathostomados
• Con mandíbula
HISTORIA EVOLUTIVA DE LOS VERTEBRADOS:
kar24239_ch03_082-127.qxd 12/21/10 7:27 PM Page 85
Adult
Larva
• Depredadores incipientes
Jaws
• Filtradores Agnathan
(grasping)
Egg
• Comportamiento filtrador Adult
Adult
• Tolerantes a ambientes de Larva
estuarios
Jaws
Agnathan
(grasping)
• Cambio a vertebrados: cambios Egg
mecánicos Adult
Vertebrate
Prevertebrate
• Faringe: bomba muscular Muscular pump
(suspension feeding)
• Contraccion de la faringe,
expulsión del agua
Gnathostome
Vertebrate
Prevertebrate
Muscular pump
• Maquinaria ciliar insuficiente (suspension feeding)
• Contraccion de la faringe,
expulsión del agua
Gnathostome
Vertebrate
Prevertebrate
Muscular pump
• Maquinaria ciliar insuficiente (suspension feeding)
complejos Adult
Larva
Egg
Vertebrate
Prevertebrate
Muscular pump
• Incrementa la resistencia (suspension feeding)
• Aparición de mandíbulas:
expanción faringea y succión mas
fuerte Cilia-mucus
(suspension feeding)
• Corporal
• Contenido de oxígeno
• Densidad
• Regulación de la temperatura
• Corporal
• Corporal
HISTORIA EVOLUTIVA DE LOS
VERTEBRADOS: DEL AGUA A LA TIERRA
• Contenido de oxígeno
• Densidad
• Ambiente terrestre