Eimeria


Eimeria is a genus of apicomplexan parasites that includes various species capable of causing the disease coccidiosis in animals such as cattle, poultry, dogs, cats, and smaller ruminants including sheep and goats. Eimeria species are considered to be monoxenous because the life cycle is completed within a single host, and stenoxenous because they tend to be host specific, although a number of exceptions have been identified. Species of this genus infect a wide variety of hosts. Thirty-one species are known to occur in bats, two in turtles, and 130 named species infect fish. Two species infect seals. Five species infect llamas and alpacas: E. alpacae, E. ivitaensis, E. lamae, E. macusaniensis, and E. punonensis. A number of species infect rodents, including E. couesii, E. kinsellai, E. palustris, E. ojastii and E. oryzomysi. Others infect poultry, rabbits and cattle. For full species list, see below.
The most prevalent species of Eimeria that cause coccidiosis in cattle are E. bovis, E. zuernii, and E. auburnensis. In a young, susceptible calf it is estimated that as few as 50,000 infective oocysts can cause severe disease. Eimeria infections are particularly damaging to the poultry industry and costs the United States more than $1.5 billion in annual loses. The most economically important species among poultry are E. tenella, E. acervulina, and E. maxima. The oocysts of what was later called Eimeria steidai were first seen by the pioneering Dutch microscopist Antonie van Leeuwenhoek in the bile of a rabbit in 1674. The genus is named after the German zoologist Theodor Eimer.

Life cycle

The Eimeria life cycle has an exogenous phase, during which the oocysts are excreted into the environment, and an endogenous phase, where parasite development occurs in the host intestine. During the endogenous phase, several rounds of asexual reproduction, or schizogony take place, after which the sexual differentiation of gametes and fertilisation occurs. Parasite transmission occurs via the oral-fecal route. Infections are common in farming environments where many animals are confined in a small space.

Oocysts

There are two forms of oocyst: sporulated or late oocyst, and unsporulated or early oocyst. An infected host releases oocysts into the environment in their unsporulated form. These contain a multi-layered cell wall making them highly resistant to environmental pressures. Once released, the unsporulated oocysts undergo meiosis upon contact with oxygen and moisture. This process is known as sporulation and the oocysts take approximately 2 to 7 days to become infectious. The sporulated oocyst is said to be tetrasporic meaning it contains four sporocysts, while each sporocyst is dizoic, i.e. it contains two sporozoites.
Once ingested, the oocysts undergo a process called excystation, whereby thousands of sporozoites are released into lumen of the intestine. In the case of E. tenella, this process is thought to occur due to the combination of enzymatic degradation and mechanical abrasion of the oocyst wall in the chicken's gizzard.

Sporozoites

The motile sporozoites invade the enterocytes of small intestine, and migrate to their respective sites of development. Invasion is mediated through specialised membrane-bound structures on the surface of the parasite that release secretions. This results in the recognition of, and attachment to host cell receptors. This process is known as gliding motility, which is conserved across all species of Apicomplexa. Membrane glyconjugates have been proposed as potential host cell receptors for Eimeria species. After invasion, the sporozoites develop into trophozoites, then into schizonts, where they undergo several rounds of asexual reproduction. This results in many nuclei developing within the schizont. Each nucleus develops into a merozoite.
Invasion requires the formation of a moving junction between parasite and host cell membranes. In E. tenella, this involves parasite micronemes and rhoptry proteins including RON2, RON5 and AMA-2. It is unlikely that the host cell is completely passive in the invasion process, although evidence of host physical forces that assist in mediating parasite entry remains controversial.

Merozoites

When schizonts rupture, merozoites are released, which either go on to re-infect more enterocytes or develop into either male or female gametes via the process of gametogenesis. These gametes fuse to form an oocyst, which is then released in its non-infectious, unsporulated form through the faeces of the host.
Merozoite invasion also requires the formation of a moving junction, however the proteins involved in this process differs from those on sporozoites. Rhoptry proteins AMA-1 and RON4 are found exclusively on merozoites. There is also a greater diversity of variant surface antigens found on the surface of merozoites. It is hypothesised that this may be due to the fact that merozoites are short-lived and a greater antigen repertoire would permit faster binding and invasion.

Taxonomy

The Eimeria lie within the family Eimeriidae. Eimeria accounts for close to 75% of the species within this family, and it is the most specious of the genera of the Apicomplexa with 1,700 described species.
Attempts to subdivide this large taxonomic unit into separate genera have been made. The classification of eimeriid coccidian was largely based on morphological and life cycle details. More recently, classification has been done using rDNA and mitochondrial genes, which indicate Eimeria may be paraphyletic to Isospora and Cyclospora
Eimeria: These species are tetrasporocystic with dizoic, nonbivalved sporocysts with or without Stieda bodies. This new genus retains the majority of the species.
Goussia : These species are tetrasporocystic, dizoic, lack Stieda bodies, and have sporocyst walls consisting of two valves joined by a longitudinal suture. This genus contains about 20 species.
Crystallospora : The species in this genus is tetrasporocystic and dizoic, and have dodecahedral sporocysts composed of two hexagonal, pyrimidal valves joined at their bases by a suture. This genus contains a single species, Crystallospora cristalloides
Epieimeria: The species in this genus are tetrasporocystic, dizoic, possess Stieda bodies, and undergo merogony and gametogony on the lumenal surface of the intestinal tract. Three species are in this genus.

Species identification

Methods for species identification are varied and among others, include isozyme analysis, the use of rRNA and rDNA probes, DNA assays and recombinant DNA techniques. PCR has proven most useful for outbreak surveillance. Prior to these methods, species identification was based on phenotypic characteristics such as the site of parasite development, the oocyst structure, the host species, cross immunity and the presence of lesions. Out of these, comparing oocyst structures was the most commonly used method.

Genomics

A whole genome sequencing project is in progress with chosen species, Eimeria tenella. The genome is about 60 megabases in size and has a GC-content around 53%. The 14 chromosomes range in size from 1 to > 6 megabases. Since 2013, the sequencing and annotation of a further six avian Eimeria species genomes is in progress.

Pathology and symptoms

Coccidiosis typically results in diarrhoea, weight loss and dehydration. A combination of these factors may result in poor growth and death of the animal, particularly amongst young. Other clinical signs include lethargy, depression, and reduced normal grooming behaviour. Diarrhoea may be bloody due to intestinal epithelium dying off when a large number of oocysts and merozoites burst out of the cells.
The severity of the disease is directly dependent on the number of infective Eimeria oocysts that are ingested. The pathogenesis of infection varies from mild to severe, and is largely dependent on the magnitude of infection. In light infections, the damage to the gut might only be minimal and be rapidly repaired as cells are rapidly replaced by the body. However, in heavy infections, it may only take two weeks for many intestinal epithelial cells to be infected with either Eimeria meronts or gametocytes. These cause the epithelial cells to burst, which causes significant damage to the intestine epithelial layer, resulting in the release of blood, fluid, and electrolytes into the intestine.

Prevention and treatment

Good animal husbandry practices and prophylactic application of anticoccidial drugs that target different stages of the parasite lifecycle, such as sulfonamides, ionophores and toltrazuril, are the preferred methods of disease prevention, particularly in the poultry industry.
The following drugs can be used for treatment of coccidiosis in cattle: amprolium, sulfaquinoxaline, and sulfamethazine. However, it is often more effective to prevent this disease in cattle, which can be aided by the products lasalocid, decoquinate, and monensin.
There is a growing problem of drug resistance, as well as possible drug residues in the meat once the animal is butchered. As a result, other avenues of control are being explored, particularly vaccine development, although several live attenuated vaccines have been in use since the 1950s. So far, the best practice is to vaccinate the chicks once they hatch from the egg so they are immune for life.
Infection with Eimeria results in life-long immunity to that particular parasite species, but does not give cross protection against other species. For these reasons, vaccines for control seem promising, of which live attenuated vaccines are most effective. However, the search for highly immunogenic antigens and overcoming antigenic variation of the parasites remains a challenge. Immunity to the parasite varies depending on parasite and host species, as well as the site of invasion. CD4+ T cells and interferon gamma are crucial components of natural immunity to infection. Humoral immunity is thought to play little role in protection, and is most likely mediated through secretory IgA antibodies.

Host-parasite relations

Fish

Eimeria aurati - goldfish

Eimeria baueri - crucian carp

Eimeria lepidosirenis - South American lungfish

Eimeria leucisci - common barbel

Eimeria rutili - European chub, Iberian nase

Eimeria vanasi - blue tilapia

Reptiles

Eimeria amphisbaeniarum - Mann's worm lizard

Eimeria witcheri - Mann's worm lizard

Eimeria yemenensae - rock agama

Birds

Eimeria acervulina - chicken
Eimeria adenoeides - turkey
Eimeria brunetti - chicken
Eimeria colchici - common pheasant

Eimeria curvata - ruddy ground dove, scaled dove

Eimeria dispersa - turkey, bobwhite quail

Eimeria duodenalis - pheasant

Eimeria fraterculae - Atlantic puffin

Eimeria gallopavonis - turkey
Eimeria innocua - turkey
Eimeria praecox - chicken
Eimeria maxima - chicken
Eimeria meleagridis - turkey

Eimeria meleagrimitis - turkey

Eimeria mitis - chicken

Eimeria necatrix - chicken

Eimeria phasiani - pheasant

Eimeria procera - grey partridges

Eimeria purpureicephali - red-capped parrot

Eimeria tenella - chicken

Mammals

Eimeria ahsata - goat, sheep

Eimeria alabamensis - cattle

Eimeria alijevi - goat

Eimeria aspheronica - goat

Eimeria arloingi - goat

Eimeria arundeli - common wombat

Eimeria bakuensis - sheep

Eimeria bovis - cattle

Eimeria cameli - camels

Eimeria caprina - goat

Eimeria caprovina - goat

Eimeria christenseni - goat

Eimeria clethrionomyis - red-backed vole

Eimeria coecicola - rabbit

Eimeria contorta - mouse

Eimeria couesii - rice rat

Eimeria crandallis - sheep

Eimeria dammahensis - scimitar-homed oryx

Eimeria dowleri - eastern red bat

Eimeria exigua - rabbit

Eimeria falciformis - mouse

Eimeria farasanii - mountain gazelle

Eimeria ferrisi - mouse

Eimeria flavescens - rabbit

Eimeria gallatii - red-backed vole

Eimeria granulosa - goat

Eimeria hirci - goat

Eimeria intestinalis - rabbit

Eimeria irresidua - rabbit

Eimeria intricata - goat

Eimeria jolchijevi - goat

Eimeria krijgsmanni - mouse

Eimeria larimerensis - Uinta ground squirrel

Eimeria macusaniensis - llamas, guanacos, alpacas, vicunas

Eimeria magna - rabbit

Eimeria marconii - red-backed vole

Eimeria media - rabbit

Eimeria melanuri - garden dormouse

Eimeria myoxi - garden dormouse

Eimeria nagpurensis - rabbit

Eimeria nieschulzi - brown rat

Eimeria ninakohlyakimovae - goat

Eimeria ovinoidalis - sheep

Eimeria pallida - goat

Eimeria palustris - marsh rice rat

Eimeria papillata - mouse

Eimeria perforans - rabbit

Eimeria phocae - Sable Island harbour seals

Eimeria pileata - red-backed vole

Eimeria pipistrellus - Kuhl's pipistrelle

Eimeria piriformis - rabbit

Eimeria prionotemni - Bennett's wallaby

Eimeria procyonis - raccoon

Eimeria punctata - goat

Eimeria roobroucki - rabbit

Eimeria saudiensis - Arabian oryx

Eimeria sealanderi - eastern red bat

Eimeria separata - mouse, rat

Eimeria stiedae - rabbit

Eimeria ursini - southern hairy nosed wombat

Eimeria vermiformis - mice

Eimeria weybridgensis - sheep

Eimeria wobati - southern hairy-nosed wombat

Eimeria zuernii - cattle

List of species

Eimeria abramovi

Eimeria acervulina

Eimeria adenoides

Eimeria ahsata

Eimeria airculensis

Eimeria alabamensis

Eimeria albigulae

Eimeria alijevi

Eimeria alpacae

Eimeria amphisbaeniarum

Eimeria anatis

Eimeria anguillae

Eimeria ankarensis

Eimeria anseris

Eimeria arizonensis

Eimeria arabukosokokensis

Eimeria arnyi

Eimeria arundeli

Eimeria anseris

Eimeria arkhari

Eimeria arloingi

Eimeria aspheronica

Eimeria auburnensis

Eimeria augusta

Eimeria aurati

Eimeria aythyae

Eimeria azerbaidschanica

Eimeria bactriani

Eimeria bakuensis

Eimeria bareillyi

Eimeria baueri

Eimeria battakhi

Eimeria beckeri

Eimeria beecheyi

Eimeria berkinbaevi

Eimeria brinkmanni

Eimeria bombaynsis

Eimeria bonasae

Eimeria boschadis

Eimeria bovis

Eimeria brantae

Eimeria brasiliensis

Eimeria brevoortiana

Eimeria brinkmanni

Eimeria brunetti

Eimeria bucephalae

Eimeria bufomarini

Eimeria bukidnonensis
Eimeria burdai

Eimeria callospermophili

Eimeria californicenis

Eimeria cameli

Eimeria canadensis

Eimeria canis

Eimeria caprina

Eimeria caprovina

Eimeria carinii

Eimeria carpelli

Eimeria catostomi

Eimeria catronensis

Eimeria caviae

Eimeria cerdonis

Eimeria citelli

Eimeria chelydrae

Eimeria christenseni

Eimeria clarkei

Eimeria clethrionomyis

Eimeria coecicola

Eimeria colchici

Eimeria columbae

Eimeria columbarum

Eimeria contorta

Eimeria coturnicus

Eimeria couesii

Eimeria crandallis

Eimeria crassa

Eimeria curvata

Eimeria cylindrica

Eimeria cynomysis

Eimeria cyprini

Eimeria dammahensis

Eimeria danailovi

Eimeria danielle

Eimeria debliecki

Eimeria deserticola

Eimeria dispersa

Eimeria dolichotis

Eimeria dromedarii

Eimeria duszynskii

Eimeria ellipsoidalis

Eimeria elongata

Eimeria etheostomae

Eimeria eutamiae

Eimeria exigua

Eimeria falciformis

Eimeria fanthami
Eimeria farasanii

Eimeria farra

Eimeria faurei

Eimeria fernandoae

Eimeria ferrisi

Eimeria filamentifera

Eimeria franklinii

Eimeria fraterculae

Eimeria freemani

Eimeria fulva

Eimeria funduli

Eimeria gallatii

Eimeria gallopavonis

Eimeria gasterostei

Eimeria gilruthi

Eimeria glenorensis

Eimeria gokaki

Eimeria gonzalei

Eimeria gorakhpuri

Eimeria granulosa

Eimeria grenieri

Eimeria guevarai

Eimeria hagani

Eimeria haneki

Eimeria hasei

Eimeria hawkinsi

Eimeria hermani

Eimeria hindlei

Eimeria hirci

Eimeria hoffmani

Eimeria hoffmeisteri

Eimeria hybognathi

Eimeria ictaluri

Eimeria illinoisensis

Eimeria innocua

Eimeria intestinalis

Eimeria intricata

Eimeria iroquoina

Eimeria irresidua

Eimeria ivitaensis

Eimeria judoviciani

Eimeria kinsellai

Eimeria koganae

Eimeria kotlani

Eimeria krijgsmanni

Eimeria krylovi

Eimeria kunmingensis

Eimeria lagopodi
Eimeria lamae

Eimeria langebarteli

Eimeria larimerensis

Eimeria lateralis

Eimeria laureleus

Eimeria lepidosirenis

Eimeria leucisci

Eimeria ludoviciani

Eimeria macusaniensis

Eimeria magnalabia

Eimeria marconii

Eimeria maxima

Eimeria melanuri

Eimeria meleagridis

Eimeria menzbieri

Eimeria micropteri

Eimeria minasensis

Eimeria mitis

Eimeria monacis

Eimeria morainensis

Eimeria moronei

Eimeria mulardi

Eimeria muta

Eimeria myoxi

Eimeria myoxocephali

Eimeria natricis

Eimeria necatrix

Eimeria neitzi

Eimeria nieschulzi

Eimeria nigricani

Eimeria nocens

Eimeria nyroca

Eimeria ojastii

Eimeria ojibwana

Eimeria onychomysis

Eimeria oryzomysi

Eimeria oryxae

Eimeria os

Eimeria osmeri

Eimeria ovata

Eimeria ovinoidalis

Eimeria palustris

Eimeria papillata

Eimeria parvula

Eimeria pigra

Eimeria pilarensis

Eimeria pileata

Eimeria pipistrellus

Eimeria phocae
Eimeria praecox

Eimeria prionotemni

Eimeria pseudospermophili

Eimeria pulchella

Eimeria pungitii

Eimeria punonensis

Eimeria ranae

Eimeria reedi

Eimeria reichenowi

Eimeria ribarrensis

Eimeria rjupa

Eimeria rutili

Eimeria salvelini

Eimeria saitamae

Eimeria saudiensis

Eimeria separata

Eimeria schachdagica

Eimeria sevilletensis

Eimeria sinensis

Eimeria sipedon

Eimeria somateriae

Eimeria spermophili

Eimeria squali

Eimeria stiedai

Eimeria stigmosa

Eimeria striata

Eimeria subepithelialis

Eimeria surki

Eimeria tamiasciuri

Eimeria tedlai

Eimeria tenella

Eimeria truncata

Eimeria truttae

Eimeria uekii

Eimeria uniungulati

Eimeria ursini

Eimeria vilasi

Eimeria weddelli

Eimeria weybridgensis

Eimeria witcheri

Eimeria vanasi

Eimeria vermiformis

Eimeria volgensis

Eimeria wobati

Eimeria wyomingensis

Eimeria yemenensae

Eimeria yukonensis

Eimeria zuernii''