Monitoring at MacMillan Wharf, Provincetown, MA

Monitoring at MacMillan Wharf, Provincetown, MA
Marine Invasive Species (MIS) Monitoring at MacMillan Wharf, Provincetown, MA.

Saturday, June 30, 2012

Ascidian Filter Feeding Mechanism

Styela clava as the model

Ascidians are suspension feeder that filter food particles such as phytoplankton from the surrounding sea water.  They transport water through the branchial sac (pharynx) which is perforated with small, ciliated slits, the stigmata.  The ciliary tracts on each side of the stigmata create a water current that pulls water through the branchial siphon into the branchial sac, through the stigmata, and into the atrial cavity from which the water leaves the body as a stream through the atrial siphon. When the water is transported across the branchial wall, suspended particles are trapped on a mucous net that is created by the endostyle, a mucus secreting structure on the ventral wall. 

Diagram of the Ascidan Filter Feeding Process

Diagram of the ascidian filter-feeding process. From Woods Hole Oceanographic Institute Website, 2005

The feeding mechanism is similar in different ascidians regardless of the complexity of the structures. Styela clava belongs to a group of solitary ascidians with a structurally complex branchial sac that is folded.  The folding increases the internal surface area, thereby increasing the number of stigmata and filtration capacity.   A mucus secreting organ on the vental surface, the endostyle, secretes a mucus net that moves across the surface of the sac by cilia on the pharyngeal bars.  The mucus net with its retained food particles then moves toward the esophagus. Thus, the amount of food consumed is determined by the concentration of food particles in the sea water, the efficiency of particle retention, and the volume of water transported. At a structural level, the endostyle is a longitudinal ciliated groove on the ventral wall and is composed of 8 different types of cells, a single band at the base of the groove with very long cilia that extend toward the surface of the groove and 7 pairs of bands on each side of the endostyle wall.

Diagram of the Branchial Sac of S. clava in Relation to the Digestive Tract

 The digestive tract of Styela clava from the left side with the internal surface of the branchial sac exposed.   bs, branchial siphon;  bt, branchial tentancles;  en, endostyle;  bb, branchial bar;  s, stigmata; i, intestine;  as, atrial siphon;  dt, dorsal tubercle;  r, rectum;  dl, dorsal lamina;  es, esophagus;  rb, retropharyngeal band;  st, stomach.

 Cross Section through the Branchial Sac of Styela clava
Section passes through the tranverse bars and not the stigmata.  The endostyle is located on the ventral surface.  Mucus and entrapped food pass along the ciliated branchial bars towards the dorsal lamina and then towards the esophagus.  bf, branchial folds;  d, dorsal lamina;  en, endostyle;  lb, longitudinal bars;  sv, subendostyle vessel.  

View of the Internal Surface of the Branchial Basket of Styela clava
Internal surface of the branchial basket of Styela clava by scanning electron microscopy (SEM) showing branchial fold (left 1/3 of image) and direct view of rows of stigmata, major and minor trasverse folds, and longitudinal folds.

Internal View of 3 Stigmata in the Branchial Sac of Styela clava
Internal surface of the branchial sac of Styela clava by SEM showing 3 stigmata, a major transverse fold at the lower left and a minor transverse fold at the upper right.  Densely packed cilia are uniform in length. 

Section through the Stigmata and a Branchial Bar of Styela clava

Section through the stigmata and a branchial bar of Styela clava. Cilia prevent food particles from leaving the branchial cavity but allow filtered sea water to pass through.  Ciliated bars transport food-containing mucus along the longitudinal and transverse bars to the dorsal lamina and thereafter to the esophagus. The cilia on the bars actually pull the mucus net forward, whereas the stigmata cilia act only as filters. sb, stigmata bar, ve, ciliated ventral epithelium;  ue, unciliated epithelium. 

Histologic Section through the Stigmata of Styela clava
Semi-cross section through a group of about a dozen ciliated stigmata ending on the left at a branchial bar.  The ascidian had contracted and stopped feeding, so the branchial sac is not in its normally expanded, feeding configuration.  Hemotoxylin stain.  

Cross section of the Endostyle of Styela clava

  Cross-sectional diagram of the different cell types of the endostyle of Styela clava.   The cells at the base of the groove have very long, flagella-like cilia.  The remaining cells are paired with semi-alternating glandular and ciliated cells.  

Histological Section of the Endostyle of Styela clava
Cross section through the endostyle of Styela clava.  The long cilia in Zone 1 can be seen extending part-way up the depth of the groove.  Hemotoxylin stain. 

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