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Sponges and Placozoans

Chapter 12

Origin of Metazoa

Evolution of the eukaryotic cell was followed by diversification into many lineages including:
Modern protozoans  Plants  Fungi  Animals

Multicellular animals are called metazoans.

Dendrogram of Major Phyla
Echinodermata
Vertebrata Uniramia Crustacea Chelicerata Priapulida Kinorhyncha Loricifera Annelida Mollusca Bryozoa Brachiopoda Phoronida Platyhelminthes Rotifera Cnidaria Porifera Nematoda

Protochordata
Hemichordata Apicomplexa Ciliophora

Actinopoda

Euglenozoa
Rhizopoda

Granuloreticulosa

.Choanoflagellates  Choanoflagellates are solitary or colonial protozoans with a flagellum surrounded by a collar of microvilli.

Choanoflagellates  Choanoflagellates resemble sponge feeding cells (choanocytes).  Scientists are studying colony formation and cell-tocell communication in choanoflagellates in search of clues to the evolution of multicellularity. .

 Trend toward bilateral symmetry as in flatworms. Recall multiple nuclei in reproducing ciliates.  . each nucleus becomes partitioned.Syncitial Ciliate Hypothesis  Syncitial ciliate hypothesis – metazoans arose from an ancestor shared with single celled ciliates.  Later.

Syncitial Ciliate Hypothesis  Problems: In flatworm embryology nothing like cellularization occurs.  .  Implies that radial symmetry is derived.  Does not explain flagellated sperm in metazoans.

Individual cells became specialized for different functions. spherical colony of flagellated cells.  Radially symmetrical. similar to a blastula.  First proposed by Haeckel in 1874  .Colonial Flagellate Hypothesis  Colonial Flagellate Hypothesis – metazoans descended from ancestors characterized by a hollow.

Phylum Porifera.Phylum Porifera  Sponges. are multicellular heterotrophs.  They lack true tissues and organs.  They are asymmetrical.  Kingdom Animalia is monophyletic. .  Molecular evidence suggests they do share a common ancestor with other animals.

 They live in both fresh and marine waters.Phylum Porifera  Sponges are sessile animals that have a porous body and choanocytes.  Supported by a skeleton of tiny needlelike spicules and protein. .

boring. finger. .   Up to 2 meters in diameter! Encrusting. tube or vase shaped.Phylum Porifera  Sponges range in size and shape.

. including crabs.Neighbors  Many organisms. and fish live as commensals or parasites in sponges. bryozoans. mites. nudibranchs.

Skeletal Framework   The skeletal framework of a sponge may be fibrous or rigid.   . The fibrous part comes from collagen fibrils in the intercellular matrix.  Spongin Calcareous Siliceous  Rigid skeletons consist of needlelike spicules.

.Suspension Feeders  Sponges are suspension feeders capturing food particles suspended in the water that passes through their body.

 It flows past the choanocytes where food particles are collected on the choanocyte collar. .Suspension Feeders  Water flows in through incurrent pores called dermal ostia.

Protein molecules are taken in by pinocytosis.Suspension Feeders Choanocytes take in small particles by phagocytosis.  Sponges can also absorb nutrients dissolved in the water.  .

Found only in the Class Calcarea. Usually tube shaped. .Canal Systems  Asconoid – the simplest canal system. Water enters through the ostia and exit through the large osculum.     Choanocytes line the spongocoel.

The walls of the sponge are folded to form choanocyte lined canals.  Increased area for feeding. .  Class Calcarea.Canal Systems   Syconoid – tubular body and singular osculum like asconoids.

permits an increase in sponge size. Choanocytes line the walls of small chambers where they can filter all the water that flows through. .Canal Systems    Leuconoids – most complex. Most sponges.

 .  Respiration and excretion occur by diffusion in each cell.Types of Cells Absence of tissues & organs means that fundamental processes occur on the cellular level.  Mesohyl is the gelatinous matrix containing skeletal elements & amoeboid cells.

flagellated collar cells. . generate a water current through the sponge and ingest suspended food.Types of Cells  Choanocytes.

& collegen (collenocytes).Types of Cells    The choanocytes pass food particles to archaeocyte cells for digestion. spongin (spongocytes). Digestion occurs entirely within cells. Other cell types secrete spicules (sclerocytes). there is no gut. .

.Types of Cells  Pinacocytes are thin.  Almost a true tissue. flat. epithelial-type cells that cover the exterior and some interior surfaces of the sponge.

 Internal buds (gemmules) in freshwater sponges can remain dormant in times of drought.Reproduction  Sponges have remarkable regeneration capabilities. .  External buds can break off to form new sponges.  Regeneration following fragmentation is a form of asexual reproduction.

Monoecious  Gametes are derived from choanocytes or sometimes archaeocytes.Reproduction  Most sponges are hermaphrodites meaning that each individual functions as both male and female.  .

 .  Some are oviparous releasing gametes into the water. Ciliated larvae are later released. the zygote is retained and is nourished by the parent.  After fertilization.Reproduction Most sponges are viviparous.

.   Flagellated cells become choanocytes & archaeocytes. Larger cells become pinacocytes.Reproduction  Sponges in the class Calcarea and a few Demospongiae have an unusual developmental pattern where the embryo turns inside out.

syconoid. usually vase shaped. . or leuconoid in structure.  Small.Class Calcarea  Calcareous sponges (Class Calcarea) have spicules composed of calcium carbonate.  Asconoid.

Chambers appear to correspond to both syconoid and leuconoid types.Class Hexactinellida  Glass sponges (Class Hexactinellida) are mostly deep sea forms.   Hexactinellids lack a pinacoderm or gelatinous mesohyll.  Spicules are six-rayed and made of silica. .

. Collar bodies do not participate in phagocytosis – this is the function of the primary and secondary reticula.  Trabecular reticulum is largest continuous syncytial tissue known in Metazoa.Class Hexactinellida     Some advocate placing hexactinellids in a subphylum separate from other sponges.forms the chambers opening to spongocoel. Trabecular reticulum made of a fusion of archaeocyte pseudopodia . Choanoblasts are associated with flagellated chambers.

 All leuconoid. .Class Demospongiae  Class Demospongiae contains most of the sponge species. but not six-rayed. or absent.  Spicules may be bound together by spongin. mostly marine.  Spicules are siliceous.

Cladogram of Sponge Classes .

  Molecular rRNA evidence suggests a Common ancestor for choanoflagellates and metazoans. .Phylogeny and Adaptive Diversification   Sponges appeared before the Cambrian.sponges arose from choanoflagellates.  Sponges and Eumetazoa are sister groups with Porifera splitting off before radiates and placozoans.  Glass sponges expanded in the Devonian. and sponges acquire them late in development One theory . However. some corals and echinoderms also have collar cells.

  No symmetry No muscular or nervous organs  Placozoans glide over food. and absorb nutrients. secrete digestive enzymes.Phylum Placozoa  Trichoplax adhaerens is the sole species of phylum Placozoa (marine). .

Phylum Placozoa  Cell layers    Dorsal epithelium Thick ventral epithelium of monociliated cells and nonciliated gland cells.  . Space between the epithelia contain fibrous “cells” within a contractile syncytium. Dorsal epithelium represents ectoderm and ventral epithelium represents endoderm.  Grell considers it diploblastic.