Organismal Biology #22
FLATWORMS AND RELATED ANIMALS

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Flatworms and nearly all other animals from here on are bilaterally symmetrical (right and left halves are mirror images).
The front end of such animals usually forms a distinct head.
However, flatworms still have a single all-purpose cavity with only one opening.

Bilateral symmetry: Right and left halves are mirror images of one another.

Phylum Platyhelminthes (flatworms): Bilaterally symmetrical animals with a flat body; dorsal (top) and ventral (bottom) surfaces differ; no circulatory system needed because every part of body is near a surface.
Anterior (front) end differs from posterior (hind) end. Sense organs and brain are concentrated at the front end (cephalization) to form a head.
A single all-purpose gastrovascular cavity, as in coelenterates; single opening functions as both mouth and anus.
A simple, ladder-like nervous system, more concentrated at the head end.
Simple excretory tubules (flame cells whose beating cilia resemble a flickering flame).
Three germ layers: ectoderm (outer epidermis); endoderm (lining of gut); mesoderm (a loose mesenchyme in flatworms). Acoelomate (no body cavity).
Many flatworms can regenerate missing parts following injury.
  • Class Turbellaria: Mostly free-living; digestive tract and sense organs still present; mouth often in middle of ventral surface.
  • Class Trematoda: Small, parasitic worms (flukes) with small, oval bodies; digestive tract simple; mouth at anterior end.
  • Class Cestoda: Highly degenerate internal parasites (tapeworms) with greatly reduced digestive tract, nervous system, and sense organs.
Flatworms, etc.

Related phyla, also without body cavities:
  • Phylum Mesozoa: Small, marine parasites with very few cells.
  • Phylum Acoela: Small, simple, bilateral animals similar to flatworms, but with no gut or digestive tract. Formerly included in the Platyhelminthes, but many zoologists now regard them as the most primitive of bilateral animals.
  • Phylum Rhynchocoela (Nemertea): "Proboscis worms," with a long, barb-tipped proboscis (or "evert") that can be protruded as a weapon or withdrawn (inverted, like the finger of a glove) when not in use.
  • Phylum Gnathostomulida: Small worms; outer epidermal cells each have a single cilium; mouth with paired, cuticle-hardened jaws.






EVOLUTION OF BILATERAL ANIMALS

Bilateral animals above the flatworm stage evolved a complete "assembly-line"
    digestive tract running from mouth to anus. Most also evolved body cavities.
Analysis of RNA sequences allows scientists to divide bilateral animals into:
    Protostomes— bilateral animals in which early cleavages are spiral and
        determinate, and in which the mouth forms early from the blastopore.
      Protostomes are further divided into:
        Lophotrochozoa, containing the Mollusca, Annelida, Bryozoa, etc.; and
        Ecdysozoa, contining the Nematoda, Arthropoda and several smaller phyla.       Deuterostomes— bilateral animals whose early cleavages are radial and
        indeterminate, and whose mouth forms at the other end from the blastopore.



FROM THIS POINT ON, all remaining phyla share several important derived features:
  • A complete "assembly line" digestive tract (mouth to anus).
  • Some type of body cavity, either a pseudocoel (a persistent blastocoel) or a true coelom (surrounded with mesoderm throughout).

"Assembly line" digestion: Nearly all animals above the flatworm level have a complete digestive tract, with a separate entrance (mouth) and exit (anus). This allows food to be processed in stages, in the manner of an assmbly line, with different regions or organs specialized for different sequential steps or for different nutrients.

Evolution of body cavities: Fluid-filled body cavities, whatever their origin, are useful:
  • in support, as a hydrostatic skeleton
  • in burrowing, where inflation of the body cavity can swell and anchor part of the body, or else wedge forward and push sediment aside.
Because of their usefulness, body cavities have evolved many times, independently, and are often constructed differently in different phyla:
  • Some animals have a pseudocoel, lined with both endoderm and mesoderm, derived from persistence of the blastocoel cavity.
  • Other animals have a true coelom, lined with mesoderm throughout. This may be either an enterocoel, derived from outpouching of the gut (as in starfish), or a schizocoel, arising within the mesoderm by splitting (as in mammals).
Differences in the structure of the coelom are useful in distinguishing many phyla, but are a poor guide to relationships among phyla because body cavities have evolved repeatedly and independently.
Animal family tree

Evolution of Bilateria



Protostomes are bilateral animals sharing the following traits:
  • The opening to the embryonic archenteron becomes the mouth
    (protostome means "first mouth").
  • Spiral cleavage, introducing an asymmetry in the 8-celled stage; the top 4 cells are rotated clockwise or counterclockwise with respect to the lower 4 cells.
  • Determinate cleavage, meaning that the cells destined to form the front left portion of the animal lose the ability to form structres on the right or the rear.
Deuterostome animals (considered later) have the opposite traits.


Phylogeny and classification of bilateral animals: Studies of ribosomal RNA sequences show evidence that bilateral animals evolved in three large groups (the first two are protostomes):
  • Lophotrochozoa: A large group that includes annelid worms, mollusks, and bryozoa, characterized in some cases by a ciliated feeding organ called a lophophore and in other cases by a ciliated larval stage called a trochophore.
  • Ecdysozoa: A group that includes the two largest phyla, Arthropoda and Nematoda, characterized by a hard outer covering that must be shed periodically during growth, using steroid hormones (ecdysones) to control the molting process.
  • Deuterostomes, including the chordates and echinoderms.
  • REVIEW:         Study guide and vocabulary

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