In addition to the single apical flagellum surrounded by actin-filled microvilli that characterizes choanoflagellates, the internal organization of organelles in the cytoplasm is constant (Leadbeater and Thomsen, 2000). A flagellar basal body sits at the base of the apical flagellum, and a second, non-flagellar basal body rests at a right angle to the flagellar base. The nucleus occupies an apical-to-central position in the cell, and food vacuoles are positioned in the basal region of the cytoplasm (Leadbeater and Thomsen, 2000; Karpov and Leadbeater, 1998). Additionally, the cell body of many choanoflagellates is surrounded by a distinguishing extracelluar matrix or periplast. These cell coverings vary greatly in structure and composition and are used by taxonomists for classification purposes. Many choanoflagellates build complex basket-shaped "houses" called lorica, from several silica strips cemented together.The functional significance of the periplast is unknown, but in sessile organisms, it is thought to aid in attachment to the substrate. In planktonic organisms, there is speculation that the periplast increases drag, thereby counteracting the force generated by the flagellum and increasing feeding efficiency (Leadbeater and Kelly, 2001).
Choanoflagellates are either free-swimming in the water column or sessile, adhering to the substrate directly or through either the periplast or a thin pedicel (Leadbeater, 1983). Although choanoflagellates are thought to be strictly free-living and heterotrophic, a number of choanoflagellate relatives such as members of Ichthyosporea or Mesomycetozoa follow a parasitic or pathogenic lifestyle (Mendoza, 2002). The life histories of choanoflagellates are poorly understood. Many species are thought to be solitary; however coloniality seems to have arisen independently several times within the group and colonial species retain a solitary stage (Leadbeater, 1983).
Choanoflagellates grow vegetatively, with many species undergoing longitudinal fission (Karpov and Leadbeater, 1998); however, the reproductive life cycle of choanoflagellates remains to be elucidated. Currently, it is unclear whether there is a sexual phase to the choanoflagellate life cycle. Interestingly, some choanoflagellates can undergo encystment, which involves the retraction of the flagellum and collar and encasement in an electron dense fibrillar wall. Upon transfer to fresh media excystment occurs, though it remains to be directly observed (Leadbeater and Karpov, 2000). Further examination of the choanoflagellate life cycle will be informative about mechanisms of colony formation and attributes present before the transition to multicellularity.
Choanoflagellates resemble the individual choanocyte cells of sponges:
There are over 125 extant species of choanoflagellates. distributed globally in marine, brackish and freshwater environments from the Arctic to the tropics, occupying both pelagic and benthic zones. Although most sampling of choanoflagellates has occurred between 0 m and 25 m, they have been recovered from as deep as 300 m in open water (Thomsen, 1982) and 100 m under Antarctic ice sheets (Buck and Garrison, 1988). Many species are hypothesized to be cosmopolitan on a global scale [e.g., Diaphanoeca grandis has been reported from North America, Europe and Australia (OBIS)], while other species are reported to have restricted regional distributions (Thomsen, et al., 1991). Co-distributed choanoflagellate species can occupy quite different microenvironments, but in general, the factors that influence the distribution and dispersion of choanoflagellates remain to be elucidated.
The choanoflagellates feed on bacteria and link otherwise inaccessible forms of carbon (since it is so small) to organisms higher in the trophic chain. Even today they are important in the carbon cycle and microbial food web.
The choanocytes (also known as "collared cells") of sponges (considered the most basal metazoa) have the same basic structure as choanoflagellates. Collared cells are occasionally found in a few other animal groups, such as flatworms.
Genome sequencing shows that among living organisms, the choanoflagellates are most closely related to animals.
The last common ancestor of animals and choanoflagellates was unicellular, perhaps forming simple colonies; in contrast, the last common ancestor of all animals was a relatively complex multicellular organism, with differentiated tissues, a definite "body plan", and complex embryonic development (including gastrulation). The timing of the splitting of these lineages is difficult to constrain, but was probably in the late Precambrian, >.
Members of the order Choanoflagellida are divided into three families based upon the composition and structure of their periplast: Codosigidae, Salpingoecidae and Acanthoecidae. Members of the family Codosigidae appear to lack a periplast when examined by light microscopy, but may have a fine outer coat visible only by electron microscopy. The family Salpingoecidae consists of species whose cells are encased in a firm theca that is visible by both light and electron microscopy. The theca is a secreted covering predominately comprised of cellulose or other polysaccharides (Adl, et al., 2005). The third family of choanoflagellates, the Acanthoecidae, contains species whose cells rest in a basket-like lorica composed of siliceous ribs or “costae” (Leadbeater and Kelly, 2001; Leadbeater and Thomsen, 2000). Discussion of Phylogenetic Relationships
The choanoflagellate families and their relationships to each other have not been tested within a phylogenetic framework. The reconstruction of the internal relationships of choanoflagellates is central to the goal of polarizing character evolution within the clade. Resolution of the internal relationships and character polarity within choanoflagellates will be informative about the character states that are ancestral within choanoflagellates and suggestive of the characteristics of the last unicellular ancestor of animals. Relationship of Choanoflagellates to Metazoans
Dujardin, a French biologist interested in protozoan evolution, recorded the morphological similarities of choanoflagellates and sponge choanocytes and proposed the possibility of a close relationship as early as 1841 (Leadbeater and Kelly, 2001). Over the past decade, this hypothesized relationship between choanoflagellates and animals has been upheld by independent analyses of multiple unlinked sequences: 18S rDNA, nuclear protein-coding genes, and mitochondrial genomes (Steenkamp, et al., 2006; Burger, et al., 2003; Mendoza, et al., 2002; Wainright, et al., 1993). Importantly, comparisons of mitochondrial genome sequences from a choanoflagellate and three sponges confirm the placement of choanoflagellates as an outgroup to Metazoa and negate the possibility that choanoflagellates evolved from metazoans (Lavrov, et al., 2005). Finally, recent studies of genes expressed in choanoflagellates have revealed that choanoflagellates synthesize homologues of metazoan cell signaling and adhesion genes (King, 2003). Because choanoflagellates and metazoans are closely related, comparisons between the two groups promise to provide insights into the biology of their last common ancestor and the earliest events in metazoan evolution.
The genome of Monosiga brevicollis, with 41.6 million base pairs, is similar in size to filamentous fungi and other free-living unicellular eukaryotes, but far smaller than that of typical animals.
The Chrysophyte Genera Synuropsis Schiller, Volvochrysis Schiller, Synochromonas Korshikov, Pseudosynura Kisselew, Pseudosyncrypta Kisselew, Chrysomoron Skuja, and Syncrypta Ehrenberg.(Statistical Data Included)
Apr 01, 2001; RUFUS H. THOMPSON  A revision of the chrysophyte genus Synuropsis Schiller is proposed and its taxonomic status is validated....