ChordataI Revised 1

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    Phylum Chordata

    The chordates are a group of particularinterest to us as we belong to it, beingmembers of the subphylum Vertebrata.

    The chordates include all of thevertebrates (fish, amphibians, reptiles,

    mammals and birds), but also two non-vertebrate subphyla: the Urochordata andthe Cephalochordata.

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    Phylum Chordata

    The chordates were in the 19thcenturyconsidered to have been derived fromprotostome ancestors (the annelid, mollusc,arthropod group).

    However, a better understanding of embryologyshows that chordates are deuterostomes and

    the invertebrates they are most closely related toare the Echinodermata (sea stars, sand dollars,sea urchins) and the Hemichordata (acornworms).

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    Important differences in development

    between protostomes and deuterostomes

    The differences in development that distinguish

    the protostomes and deuterostomes include:

    Whether cleavage of cells in the early zygote is spiral

    or radial. Whether or not, if the early blastomere is separated,

    each cell can develop into a normal larva or not.

    Whether the blastopore ultimately forms the mouth or

    anus of the organism. Whether or not the organism possesses a coelom

    and how that coelom is formed.

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    Protostomes and Deuterostomes

    Protostomes include the annelids,

    mollusks, and arthropods.

    Deuterostomes include the echinoderms

    and Chordates, which includes the

    vertebrates.

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    Figure 23.02

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    Characteristics of the Chordata

    Chordates are:

    bilaterally symmetrical

    triploblastic

    have a well developed coelom

    have a complete digestive system

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    Germ layers of a triploblastic

    organism Endoderm: innermost germ layer of an embryo. Forms

    the gut, liver, pancreas.

    Ectoderm: Outer layer of cells in early embryo.

    Surrounds the blastocoel. Forms outer epithelium ofbody and nervous system.

    Mesoderm: Third germ layer formed in gastrula between

    ectoderm and endoderm. The coelomforms in themesoderm. Mesoderm gives rise to connective tissue,muscle, urogenital and vascular systems andperitoneum.

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    Coeloms

    The coelom is a cavity entirely surroundedby mesoderm.

    A coelom provides a tube-within-a-tube

    arrangement which has many advantages:

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    Advantages of a coelom

    In mammals the pericardial, peritoneal, andpleural cavities are formed from the coelom.

    Allows flexibility in arranging visceral organs

    Permits greater size and complexity ofinternal organs as there is space for them togrow

    Fluid-filled coelom can act as a hydrostatic

    skeleton Cushions organs against impact

    Allows organs to grow and moveindependently of your outer body wall.

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    Five distinctive characteristics of

    the chordates Five distinctive characteristics separate the

    chordates from all other phyla: Notochord

    Single, dorsal, tubular nerve cord

    Pharyngeal pouches or slits

    Endostyle

    Postanal tail

    Not all of these characteristics are apparent inadult organisms and may appear only in theembryonic or larval stages.

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    Notochord

    Notochord: the notochord is a flexible,

    rodlike structure. It extends the length of

    the body and is an anchor point for

    muscles.

    The notochord bends without shortening

    so it permits the animal to undulate.

    Figure 23 01

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    Figure 23.01

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    Notochord

    In nonvertebrates and the jawless

    vertebrates the notochord is present

    throughout life.

    However, in the jawed vertebrates it is

    replaced by the vertebral column; the

    remnants of the notochord being found in

    the intervertebral disks.

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    Single, dorsal, tubular nerve

    cord In most invertebrates the nerve cord, if present,

    is ventral to the gut.

    In chordates, in contrast, the nerve cord is dorsalto the gut and notochord. The nerve cordpasses through the neural arches of thevertebrae, which protect it.

    The nerve cord is enlarged in vertebrates into abrain, which is surrounded by a bony orcartilaginous cranium.

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    Pharyngeal pouches and slits

    Pharyngeal slits occur in aquatic chordates andlead from the pharyngeal cavity to the outside.

    The pharyngeal slits are used as a filter feedingdevice in protochordates (i.e., Urochordata(Tunicates)) and Cephalochordata (lanceletse.g. Amphioxus).

    Water containing food is drawn in through themouth by cilia and exits via the pharyngeal slitswhere the particles are trapped in mucus.

    Figure 23 09b

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    Figure 23.09b

    Amphioxus

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    Pharyngeal pouches and slits

    In vertebrates the pharyngeal arches have

    been modified into gills by the addition of a

    rich blood supply and thin gas permeable

    walls.

    The contraction of muscles in the pharynx

    drive water through the gills.

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    Pharyngeal pouches and slits

    In amniotes an opening may not form and

    rather than slits only grooves called

    pharyngeal pouches develop.

    In tetrapods these pouches give rise

    during development to a variety of

    structures including the middle ear cavity,

    eustachian tube, and tonsils.

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    Endostyle or thyroid gland

    The endostyle is found in protochordates

    and in lamprey larvae. It is located on the

    floor of the pharynx and secretes mucus,

    which is used to trap particles.

    The endostyle works with the pharyngeal

    slits in filter feeding.

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    Endostyle or thyroid gland

    Some cells in the endostyle secrete

    iodinated proteins and are homologous

    with the iodinated-hormone secreting

    thyroid gland, which is found in adultlampreys and vertebrates.

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    Postanal tail

    The postanal tail, some musculataure and thenotochord enable larval tunicates andamphioxus to swim.

    The postanal tail evolved to allow organisms toswim and its efficiency has been enhanced bythe addition of fins. The postanal tail is present

    only in vestigial form in humans (the coccyx)although tails as a whole are widespread amongvertebrates.

    Figure 23.09b

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    Amphioxus

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    Classification of the Chordata

    There are three subphyla in the Chordata:

    Subphylum Urochordata: tunicates

    Subphylum Cephalochordata: lancelets

    Subphylum Vertebrata: fish, amphibians,

    reptiles, birds, mammals, etc.

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    Subphylum Urochordata

    The Urochordata (tunicates named for the

    tough tunic that surrounds the adult) look like

    most unpromising candidates to be chordates

    and relatives of the vertebrates.

    The largest group, the ascidians or sea squirts

    (Class Ascidiacea) as adults are marine, sessile,

    filter feeding organisms that live either solitarily

    or in colonies.

    http://www.habitas.org.uk/marinelife/photo.asp?item=cioint
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    Ciona intestinalis

    (a solitary sea squirt)

    0147.jpg

    http://www.habitas.org.uk/marinelife/photo.asp?item=ciointhttp://www.habitas.org.uk/marinelife/photo.asp?item=ciointhttp://www.habitas.org.uk/marinelife/photo.asp?item=cioint
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    http://www.habitas.org.uk/marinelife/photo.asp?item=synpul
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    Synoicum pulmonaria a colonial sea squirt

    http://www.habitas.org.uk/marinelife/photo.asp?item=synpulhttp://www.habitas.org.uk/marinelife/photo.asp?item=synpulhttp://www.habitas.org.uk/marinelife/photo.asp?item=synpulhttp://www.habitas.org.uk/marinelife/photo.asp?item=synpul
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    Ascidians

    Adult ascidians lack a notochord and there

    is only a single ganglion in place of the

    dorsal nerve cord.

    Of the five characteristics of chordates

    adults possess only two: pharyngeal gill

    slits and an endostyle, both of which theyuse in filter feeding.

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    Ascidians

    The adult sea squirt draws water in

    through an incurrent siphon and pushes it

    back out an excurrent one.

    Food particles are filtered out in the

    pharyngeal slits with mucus from the

    endostyle used to trap particles.

    Figure 23.04

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    15.4

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    http://seapeachbio.blogspot.com/2012/06/ascidian-filter-feeding-mechanism.html

    Ascidian filter feeding process. Water is drawn

    through the animal by the beating of cilia.

    Cilia lining the slits in the branchial sac or pharynx

    Branchial sac orpharynx

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    Larval Ascidian

    Even though the adult ascidian hardly

    resembles a chordate its larva does.

    Larval ascidians are very small and

    tadpole-like and possess all five chordate

    characteristics previously outlined.

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    Young larval ascidian

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    Larval Ascidian

    The larval ascidians role is to disperse andto achieve this it is free swimming.However, it has only a short larval life

    (minutes to a couple of days) and does notfeed during this time.

    Instead it searches for a place to settleand then attaches and metamorphosesinto an adult.

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    Ascidian metamorphosis

    During metamorphosis the notochord

    disappears, the nerve cord is reduced to a

    single nerve ganglion and a couple of

    nerves.

    Figure 23.06

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    15.5

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    Other Urochordate classes

    Besides the ascidians there are two otherclasses of the Urochordata: the Larvaceaand Thaliacea.

    Both are small, transparent planktonicforms. Thaliaceans are cylindrical or

    spindle shaped whereas larvaceans aretadpolelike and resemble an ascidianlarva.

    G f

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    Garstangs hypothesis of chordate

    larval evolution

    In the 1920s it was proposed that the

    vertebrates were derived from an

    ancestral ascidian that retained its

    characteristics into adulthood (the processby which juvenile characteristics are

    retained into adulthood is referred to as

    paedomorphosis).

    Figure 23.12

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    G t h th i f h d t

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    Garstangs hypothesis of chordate

    larval evolution

    Garstangs hypothesis is supported by

    embryological evidence, but more recently

    molecular analyses have suggested that

    sessile ascidians are a derived form andthat the free-living larvaceans are more

    likely to be the closest relatives of the

    chordates.

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    Subphylum Cephalochordata

    The cephalochordates are the lancelets, whichare small (3-7 cm long) laterally compressedfishlike animals that inhabit sandy sediments ofcoastal waters. They lack a distinct head and

    have no cranium.

    They are commonly referred to as Amphioxus asthis was the original genus name. There are 29species, five of which occur in North Americancoastal waters.

    Figure 23.09b

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    Amphioxus

    Amphioxus is a filter feeder.

    Water enters the mouth and then passes

    through the pharyngeal slits, where food is

    trapped in mucus. Cilia then move the

    food to the gut.

    Figure 23.09a

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    Amphioxus

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    Amphioxus

    Amphioxus is interesting because it displays the

    basic chordate characteristics in a simple and

    obvious form because of its transparency.

    Amphioxus is considered to be the closest living

    relative of the vertebrates because it shares

    several characteristics with vertebrates that

    Urochordates do not possess.

    Amphioxus characteristics

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    Amphioxus characteristics

    shared with vertebratesAs well as possessing the chordate

    characteristics, amphioxus shares a

    number of traits with vertebrates including:

    Segmented myomeres (muscle blocks)

    Dorsal and ventral aortas

    Branchial (gill) arches (blood vessels running

    over the gills).

    Characteristics of vertebrates

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    Characteristics of vertebrates

    notpossessed by AmphioxusAmphioxus however lacks several

    characteristics that biologists think the

    ancestor of vertebrates possessed. These

    include: Tripartite brain(with forebrain, midbrain and

    hindbrain) protected by a cranium

    Chambered heart Muscular gut and pharynx

    List continues on next slide

    Characteristics of vertebrates

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    Characteristics of vertebrates

    notpossessed by Amphioxus Various special sensory organs(eyes,

    chemical and pressure receptors)

    Neural crest(ectodermal cells that are found

    on the embryonic neural tube and areengaged in the formation of the cranium, tooth

    dentine, some endocrine glands and

    Schwann cells, which provide myelin

    insulation to nerve cells).