Apicomplexa

A phylum made up almost entirely of parasites. Are distinguished by their unique method of entering
host cells.

Cause a number of serious illnesses, such as

babesiosis in dogs and cattle, leucocytozoonosis in birds, and most significantly malaria in humans.

 Grouped with dinoflagellates and ciliates to make up the

Alveolata, a higher order whose most common shared characteristic is the cortical alveolae, flattened vesicle-like structures which are found just under the plasma membrane.

Unlike their Alveolate relations, however, Apicomplexans

have undergone degenerative evolution, losing all flagella and cilia.

Their unique characteristic, the product of past

endosymbiosis, is the apical complex, a group of secretory organelles, in particular the apicoplast, which enable the parasitic cells to invade the host cell.

 Algae are organisms that have plasmids, or organisms that

are derived from cells whose ancestors possessed plastids.

Until 1994, it was thought that the apicomplexa did not have
plasmids (and consequently were not covered phycology textbooks)

 Then it was shown that

a known organelle in many apicomplexa was actually a reduced colorless plastid called an apicoplast.

General Morphological Features

All members of this phylum have an infectious stage
- the sporozoite - which possess three distinct structures in an apical complex. consists of a set of spirally arranged microtubules (the
conoid), a secretory body (the rhoptry) and one or more polar rings.

Additional slender electron-dense secretory bodies

(micronemes) surrounded by one or two polar rings may also be present.

 A further group of spherical organelles are

distributed throughout the cell rather than being localized at the apical complex and are known as the dense granules.
Secretion of the dense-granule content takes place after
parasite invasion and localization within the parasitophorous vacuole and persists for several minutes.

Ultrastructural Characteristics of Apicomplexa

Polar rings Conoid Micronemes

Rhoptries
Dense granules Pellicle (i.e. inner membrane complex)

Micropores

 The apical complex is the flag trait required for

classification as Apicomplexa.

It is a set of ultrastructurally and functionally

homologous components found at the anterior end of certain stages, most notably the infective stages, displacing the nucleus and mitochondria towards the posterior end

 . The structural (cytoskeletal) components of the apical

complex are known as the conoid, polar rings and sub-pellicular microtubulles.

Three types of distinct secretory organelles are typically

present: rhoptries, micronemes and dense granules.
*Rhoptries and micronemes are always associated with the anterior end, while dense granules are often in the posterior end as well.

 The feature that unifies Apicomplexa with other alveolates

(Alveolata) is a distinct membrane complex that envelopes all infective stages.

The outermost membrane part is called plasmalema. The

inner part is composed of two membranes running parallel to each other forming the flattened inner

membrane

complex (a.k.a. alveoli).

The pellicle is interrupted at the anterior end by the polar
rings associated with the apical complex and a single or several micropores.

 They are comprised of

endosymbiontic derived organelles (mitochondrion,
apicoplast),

universal eukaryotic organelles (nucleus, endoplasmic

reticulum, Golgi apparatus, ribosomes),

and specific compartments (i.e. acidocalcisomes),

*all enclosed by a complex membranous structure termed the pellicle.

 The discovery of the apicoplast generated considerable

interest since most apicomplexans are unicellular endoparasites that cause some of the most significant tropical diseases. Malaria in humans is produced by the apicomplexan
Plasmodium. one million deaths annually

About 300 million people are infected with malaria, leading to Apicomplexans cause other serious diseases in livestock and
humans, such as cryptosporidiosis, babesiosis, theileriosis, and toxoplasmosis.

 The realization that these endoparasites were once

algae raised hopes that the apicoplast might be a drug target for two reasons. The first is that the apicoplast is essential for the survival
of Plasmodium and Toxoplasma.

The second is that drugs effective against prokaryotic

organisms might be effective against the apicoplast since all plastids originally evolved from endosymbiotic prokaryotic cyanobacteria.

 Apicomplexans are absolutely dependent on the

apicoplast, which has led to speculation that this curious organelle is a potential Achilles heel of parasites, such as Plasmodium.

 The typical apicomplexan vegetative cell (merozoite)

has an apicoplast surrounded by four membranes.

The inner two membranes are the inner and outer

plastid membranes while the outer membranes are derived from the vacuolar membrane and the plasma membrane of the endosymbiotic red alga.

 The apical

complex consists of a
polar ring and a conoid formed of
spirally coiled microtubules.

The apicomplexan has laminin polysaccharide on its surface while the host cell has a laminin receptor.

 The apicomplexan parasite attaches to the host cell with the

conoid protruding to produce a stylet that forms a tight junction between the apicomplexan parasite and host cell.

The apicomplexan cell is taken up into the host cell in the

parasitophorous vacuole.

The contents of the rhoptries and micronemes are

emptied into the space between the apicomplexan plasma membrane and the parasitophorous vacuole membrane.

 Apicomplexans have a layer

of flattened membranous

sacs or alveoli beneath the plasma membrane that comprise the subpellicular membrane complex, similar to that found in the dinoflagellates.

History
The first apicomplexan protozoan was seen by Antonie
van Leeuwenhoekc who in 1674 saw oocysts of Eimeria stiedae in the gall bladder of a rabbit.

The first member of the phylum to be named (by Dufour

in 1828) was Gregarina ovata in earwigs. Since then many more have been identified and named.

During the quarter century 1826-1850, 41 species and 6

genera of Apicomplexa were named. In the quarter century 1951-1975, 1873 new species and 83 new genera were added.

 The apicomplexa are a monophyletic group composed

almost entirely of parasitic (ie, no free-living) species.

Apicomplexa, along with ciliates and dinoflagellates,

form a higher order group known as Alveolata.

A major defining characteristic of the this group are

flattened vesicle-like structures--called cortical alveolae-which are found just underneath the plasma membrane. Formerly the apicomplexa were part of a group called sporozoa and this name is still sometimes used. There have been some suggestions to revert back to the name sporozoa

Motile structures such as flagella or pseudopods are

present only in certain gamete stages.

 Electron microscopy revealed unique ultrastructural

features among the various sporozoa which were subsequently used to redefine the groups.

A defining characteristic of the apicomplexa is a group

of organelles found at one end--called the apical end--of the organism.

This 'apical complex' includes secretory organelles known

as micronemes and rhoptries, polar rings composed of microtubules, and in some species a conoid which lies within the polar rings.

 At some point during their life cycle, members of the

apicomplexa either invade or attach to host cells. It is during this invasive (and/or motile) stage that these apical organelles are expressed as well as the subpellicular membranes, which are actually cortical alveoli.

The apical organelles play a role in interaction of the

parasite with the host cell and the subsequent invasion of the host cell. Motile forms of apicomplexa crawl along the substratum in a non-ameboid fashion known as gliding motility. Many apicomplexan species have flagellated gametes.

 The apicomplexa have complex life cycles that are

characterized by three distinct processes: sporogony, merogony and gametogony.

Although most apicomplexa exhibit this overall

general life cycle the details can vary between species. Furthermore, the terminology used to describe these various life cycle stages vary between the species.

Life Cycle

 The life cycle consists of both asexually

reproducing forms and sexual stages.

In monoxenous species all three of these processes

will be carried out in a single host and often in a single cell type or tissue. Whereas, in heteroxenous species the various processes will be carried out in different hosts and generally involve different tissues.

 Sporogony occurs immediately after a sexual phase and

consists of an asexual reproduction that culminates in the production of sporozoites. Sporozoites are an invasive form that will invade cells and develop into forms that undergo another asexual replication known as merogony. many different names depending of the species. In contrast to sporogony, in which there is generally only one round of replication, quite often there are multiple rounds of merogony.

Merogony and the resulting merozoites are known by

 In other words, the merozoites, which are also invasive

forms, can reinvade cells and initiate another round of merogony. Sometimes these multiple rounds of merogony will involve a switch in the host organism or a switch in the type of cell invaded by the parasite resulting in distinct stages of merogony. As an alternative to asexual replication merozoites can develop into gametes through a process variously called gametogony, gamogony or gametogenesis. As in other types of sexual reproduction, the gametes fuse to form a zygote which will undergo sporogony.

Ecology
Apicomplexan hosts range from humans to rodents to
dogs to horses. Distribution varies according to species; some are concentrated in tropical or subtropical areas where the specific requirements of the parasite are meant, while others may be found in parts of North America, Europe, and Asia. Most life cycles involve two hosts: the intermediate invertebrate (insect) vector and the definitive vertebrate host, although some species may also be spread from vertebrate host to vertebrate host. Currently many Apicomplexans, most famously the genus Plasmodium, are under close observation and are being researched extensively.

Plasmodium

The apicomplexa are an

extremely large and diverse group (>5000 named species). Seven species infect humans.

Babesia
Cryptosporidium Isospora

Cyclospora
Sarcocystis Toxoplasma

Plasmodium, as the

causative agent of malaria, has the greatest impact on human health. Babesia is a relatively rare zoonotic infection.

 The other five species are all classified as coccidia.

However, recent molecular data indicates that Cryptosporidium is more closely related to the gregarines than to the coccidia. The coccidia are generally considered opportunistic pathogens and are often associated with AIDS.

Several apicomplexan parasites are also important in

terms of veterinary medicine and agriculture. Most notable areBabesia and Theileria in cattle and Eimeria in poultry.

Algal Origins of the Apicomplexa

Historically the apicomplexa have been described as a
group with only parasitic forms. This and their unique apical organelles bring up questions in regards to the origin of the group. genus Copodella form a sister group with the apicomplexa. The colpodellids are predatory flagellates that feed on unicellular algae by a process called myzocytosis..

Phylogenetic analysis indicates that members or the

 Myzocytosis involves the predator (or parasite) attaching

to the prey (or host) and literally sucking out the cytoplasm of the prey cell via specialized structures. This attachment and interaction with the prey cell is mediated by organelles similar to those that the apicomplexa utilize for attachment to or invasion of host cells.

Thus the evolution of the apicomplexa likely evolved from

this myzocytoic predation to myzocytoic parasitism, as exhibited by gregarines and Cryptosporidium, to intracellular parasitism

 Other myzocytoic organisms with apicomplexa like apical

organelles include Perkinsus, parasites of oysters and clams, and Parvilucifera, a predator of dinoflagellates. These perkinsids, however, form a sister group with the dinoflagellates and not the apicomplexa. This suggests that the progenitor of dinoflagellate and apicomplexan clades may have been a predatory flagellate and that the apical organelles were retained in the apicomplexan clade, but lost in most of the dinoflagellate clade.

 The other connection between algae and the apicomplexa

is a chloroplast remnant, called the apicoplast, found in most apicomplexans. The apicoplast is likely the result of a secondary endosymbiosis of a red algae and is likely the same endosymbiotic event giving rise to the plastids of dinoflagellates.

The apicoplast is nonphotosynthetic but exhibits activities

associated with type II fatty acid biosynthesis, isoprenoid biosynthesis, and possibly heme synthesis. These pathways are essentially prokaryotic and represent excellent drug targets. A photosynthetic alveolate, Chromera velia, that appears to be the earliest branching apicomplexan has also been identified.

 Molecular studies have shown that the apicoplast and

dinoflagellate plastids originated from red algae by single endosymbiotic event that occurred relatively early in eukaryotic evolution.

General Features
Formally Apicomplexa is split into three principal
groups; the Coccidia (=coccidia , haemogregarines), Gregarina(=gregarines) and Hematozoa (=malarial parasites, piroplasms), all of which are
obligate parasites.

It has been postulated that before any morphological

characteristic is utilised as the basis of taxonomical classification, its association with that particular group of organism has to be confirmed with clade analysis.

 The lack of such support for morphological traits, and

additional errors associated with inappropriate representation of taxa and misuse of genetic analysis tools in the classification of organisms within the three different groups has complicated this field over the years.

Recently, much emphasis has been placed on identification of

clade-supported traits in an attempt to improve classification criteria.

Taxonomy - Perkins et al (2000)

Class Aconoidasida
Order Haemospororida Order Piroplasmorida

Subclass Coccidiasina
Order Agamococcidiorida Order Eucoccidiorida Order Ixorheorida Order Protococcidiorida

Class Conoidasida
Subclass Gregarinasina
Order Archigregarinorida Order Eugregarinorida
Suborder Adeleorina SuborderEimeriorina Order Neogregarinorida

Class Perkinsasida
Order Perkinsorida
Family Perkinsidae

Gregarines
The gregarines are generally parasites
of annelids, arthropods and mollusks. They are often found in the guts of their hosts but may invade the other tissues. In the typical gregarine life cycle a trophozoite develops within a host cell into a schizont. This then divides into a number of merozoites by schizogony. The merozoites are released by lysing the host cell which in turn invade other cells. At some point in the apicomplexan life cycle, gametocytess are formed. These are released by lysis of the host cells which group together. Each gametocyte forms multiple gametes. The gametes fuse with another to form oocysts. The oocysts leave the host to be taken up by a new host.

Coccidians
Coccidians are generally parasites of vertebrates. Like gregarines they are The coccidian life cycle involves merogony, gametogony and sporogony.
commonly parasites of the epithelial cells of the gut but may infect other tissues. While similar to that of the gregarines it differs in zygote formation. Some trophozoites enlarge and become macrogamete while others divide repeatedly to form microgametes (anisogamy). The microgametes are motile and must reach the macrogamete to fertilize it. The fertilized macrogamete forms a zygote which in its turn forms an oocyst which is normally released from the body. Syzygy when it occurs involves markedly anisogamous gametes. The life cycle is typically haploid with the only diploid stage occurring in the zygote which is normally short lived.

Differences between the coccidia and the gregarines

The main difference between the coccidians and the
gregarines is in the gamonts. In the coccida these are small, intracellular and without tepimerites or mucrons. In the gregarines these are large, extracellular and posses epimerites or mucrons.

A second difference between the coccidia and the

gregarines also lies in the gamonts. In the coccidia a single gamonts becomes a macrogametocyte while in the gregarines the gamonts give rise to multiple gametocytes.

Haemosporidia
The Haemosporidians have more complex life cycles that alternate
between an arthropod and a vertebrate host. The trophozoite parasitises erythrocytesor other tissues in the vertebrate host. Microgametes and macrogametes are always found in the blood. The gametes are taken up by the insect vector during a blood meal. The microgametes migrate within the gut of the insect vector and fuse with the macrogametes. The fertilized macrogamete now becomes an ookinete which penetrates the body of the vector. The ookinete then transforms into an oocyst and divides initially by meiosis and then by mitosis (haplontic life cycle) to give rise to the sporozoites. The sporozoites escape from the oocyst and migrate within the body of the vector to the salivary glands where they are injected into the new vertebrate host when the insect vector feeds again.

 Katablepharis is a heterotrophic

unicellular flagellate that lacks a plastid. that was classified with the Cryptophyceae.

Katablepharis cells have ejectisomes

However, ultrastructural studies

revealed the presence of an anterior conoid apparatus involved in phagocytosis of prey. similar to those of the apicomplexans and it is likely that Katablepharis should be classified as an apicomplexan.

The conoid apparatus is very

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