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''Gregarina garnhami'' is a
eukaryotic Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
unicellular organism A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and ...
belonging to the
Apicomplexa The Apicomplexa (also called Apicomplexia) are a large phylum of parasitic alveolates. Most of them possess a unique form of organelle that comprises a type of non-photosynthetic plastid called an apicoplast, and an apical complex structure. T ...
described in 1956 by Canning as a
parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
found in several
locust Locusts (derived from the Vulgar Latin ''locusta'', meaning grasshopper) are various species of short-horned grasshoppers in the family Acrididae that have a swarming phase. These insects are usually solitary, but under certain circumstanc ...
s, such as the
desert locust The desert locust (''Schistocerca gregaria'') is a species of locust, a periodically swarming, short-horned grasshopper in the family Acrididae. They are found primarily in the deserts and dry areas of northern and eastern Africa, Arabia, and sou ...
, African migratory locust, and Egyptian locust. Especially, the desert locust is the host for this species, as up to 100% of animals can become infected. An estimated thousands of different species of gregarines can be in
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s and 99% of these gregarines still need to be described. Each insect is said to host multiple species. A remarkable feature of ''G. garnhami'' is its
autofluorescence Autofluorescence is the natural emission of light by biological structures such as mitochondria and lysosomes when they have absorbed light, and is used to distinguish the light originating from artificially added fluorescent markers (fluorophores) ...
.


Taxonomy

''Gregarina garnhami'' was considered by Lipa et al. in 1996 to be synonymous with ''Gregarina acridiorum'' (Léger, 1893), a parasite of several orthopteran species including ''Locusta migratoria''. Indeed, ''G. acridiorum'' and ''G. garnhami'' share common morphological and behavioral characteristics, such as their development in the midgut of their hosts, a small globular epimerite, stout bodied gamonts, and barrel-shaped (or dolioform) oocysts. In 2021, an integrative taxonomy study, using morphological and molecular characters, concluded that the two species were distinct and thus confirmed the validity of ''G. garnhami''. However, phenotypic plasticity was clearly observed in the case of ''G. garnhami'': the morphology of its
trophozoite A trophozoite (G. ''trope'', nourishment + ''zoon'', animal) is the activated, feeding stage in the life cycle of certain protozoa such as malaria-causing ''Plasmodium falciparum'' and those of the '' Giardia'' group. (The complement of the trophozo ...
s, gamonts and syzygies varied according to the geographical location of ''S. gregaria'' and the subspecies infected. Text was copied from this source, which is available under
Creative Commons Attribution 4.0 International License


Cell structure

left, Structure of a septate (right) and aseptate (left) eugregarine ''Gregarina garnhami'' is a
gregarine The gregarines are a group of Apicomplexan alveolates, classified as the Gregarinasina or Gregarinia. The large (roughly half a millimeter) parasites inhabit the intestines of many invertebrates. They are not found in any vertebrates. However, gr ...
that belongs to the septate eugregarines, meaning its cell is separated into parts. In ''G. garnhami'', three parts can be seen: epimerite, protomerite, and deutomerite, but their visibility depends on the lifestage of the organism. A characteristic of gregarines is the typical construction of the pellicula that is formed by a
cell membrane The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment ( ...
and two cytoplasmic membranes (the latter is often referred to as the inner membrane complex, IMC or alveoli). The membranes' proximity to one another often makes them difficult to distinguish. This structure is often also referred to as the trilayered structure. Beneath the inner membrane, a basal (internal) lamina can be seen, which separates the foldings around the cell from the rest of the cell. These foldings form the outer surface of'' G. garnhami'', and hundreds of these can be observed at the surface enlarging the surface of the cell. The
cytoplasm In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. The ...
of the cell (in the epimerite, protomerite, and deutomerite) is divided in two zones: ectoplasm and
endoplasm Endoplasm generally refers to the inner (often granulated), dense part of a cell's cytoplasm. This is opposed to the ectoplasm which is the outer (non-granulated) layer of the cytoplasm, which is typically watery and immediately adjacent to the ...
. The ectoplasm is clear and does not contain much granular material. In the ectoplasm, the
microtubule Microtubules are polymers of tubulin that form part of the cytoskeleton and provide structure and shape to eukaryotic cells. Microtubules can be as long as 50 micrometres, as wide as 23 to 27  nm and have an inner diameter between 11 an ...
s can be found. The endoplasm is less transparent and contains paraglycogen giving the cells a brown-yellowish color when viewing the cells with a light miscrope.


Life cycle

The lifecycle consists of several stadia: gametocyst -
oocyst Apicomplexans, a group of intracellular parasites, have life cycle stages that allow them to survive the wide variety of environments they are exposed to during their complex life cycle. Each stage in the life cycle of an apicomplexan organism i ...
-
sporozoite Apicomplexans, a group of intracellular parasites, have life cycle stages that allow them to survive the wide variety of environments they are exposed to during their complex life cycle. Each stage in the life cycle of an apicomplexan organism i ...
trophozoite A trophozoite (G. ''trope'', nourishment + ''zoon'', animal) is the activated, feeding stage in the life cycle of certain protozoa such as malaria-causing ''Plasmodium falciparum'' and those of the '' Giardia'' group. (The complement of the trophozo ...
(chepaline) - gamont (gametocyte, sporadin) – gametocyst – oocyst. The gametocyst and oocysts are the cell structures that can survive outside the host-organism and infect other insects. The cycle starts with the oocysts that leave the body with the feces and are left on plant-material. When other locust than eat the plant with oocysts they will burst open (excystation) under the influence of the digestion of the locust. Eight sporozoites will then be released inside the digestive system of the locust.
Orthoptera Orthoptera () is an order of insects that comprises the grasshoppers, locusts, and crickets, including closely related insects, such as the bush crickets or katydids and wētā. The order is subdivided into two suborders: Caelifera – grassho ...
regurgitate
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products. A ...
s from the foregut and together with secretions of the salivary glands these enzymes can break down the oocyst cell wall and thus enable the freeing of the sporozoits. The free sporozoits than pass through the peritrophic membrane surrounding the midgut. Once in the ectoperitrophic space they can attach themselves to the epithelial cells of the caeca and midgut. Once attached to a epithelial cell, it grows vegetatively and becomes a trophozoite (also called cephalin or chepalont). After 48 hours, a cell with two structures can be seen: the epimerite, attaching to the host cell and the second part (back) of the cell. After a while, a septum is formed, creating a clear protomerite and deuteromerite. After about eight days the Trophozoite will release itself from the host cell. After release from the epithelial cell, a trophozoit associates with a second one and forms a gamont by forming a circle and fusing together ( syzygy). Once no distinction between the two is visible, zygotes are formed. The zygotes are the only diploid lifestage in the lifecycle of ''G. garnhami''. A cyst (oocyst) eventually is formed and the nucleus goes through a
meiotic Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately res ...
and
mitotic In cell biology, mitosis () is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is maintai ...
division. In the end, eight haploid sporozoits are formed, ready to be released into a new host.


Relationship with their host

Their relationship with their hosts has not been deciphered in full detail, but ''G. garnhami'' is generally regarded as a commensal organism that does not harm its host. A typical characteristic is that most eugregarines only have one host during their development. This relationship is generally described as host-specific, but this can also be due to the lack of research on this topic. This host-specificity is set to take place at the level of a specific family of hosts in the case of
Orthoptera Orthoptera () is an order of insects that comprises the grasshoppers, locusts, and crickets, including closely related insects, such as the bush crickets or katydids and wētā. The order is subdivided into two suborders: Caelifera – grassho ...
hosts. Although some studies mention more hosts per species of gregarine. Lifestage specificity has been shown and this is mainly important for holometabolic insects. Due to a lack of genetic information on gregarines it is however difficult to study the host-specificity. The identification of gregarines is often based on external characteristics, but they are very similar making identification difficult. In the past (and as of 2020) identification of gregarines was often based on the host, but this seems not to be an advisable method. ''Gregarina garnhami'' is mainly found in the caeca and midgut of the desert locust where it attaches itself to the epithelial cells in order to feed. The uptake of food happens through the epimerite, attached to the host cell, after a while also the developed protomerite and deutomerite can take up nutrients. The uptake of nutrients happens through
osmosis Osmosis (, ) is the spontaneous net movement or diffusion of solvent molecules through a selectively-permeable membrane from a region of high water potential (region of lower solute concentration) to a region of low water potential (region of ...
and
pinocytosis In cellular biology, pinocytosis, otherwise known as fluid endocytosis and bulk-phase pinocytosis, is a mode of endocytosis in which small molecules dissolved in extracellular fluid are brought into the cell through an invagination of the cell me ...
(formation of a
Cytostome A cytostome (from ''cyto-'', cell and ''stome-'', mouth) or cell mouth is a part of a cell specialized for phagocytosis, usually in the form of a microtubule-supported funnel or groove. Food is directed into the cytostome, and sealed into vacuole ...
). The nutrients are then stored as paraglycogen or fat. The hundreds of foldings of the cell surface give eugregarines a substantial enlargement of the cell-surface in order to take up more nutrients. More details need to be studied in regard of the nutrient uptake by gregarines, however, it is clear that they do not pierce the host cell to take up the cytoplasm. An apical complex is absent in the trophozoit stage in most eugregarines while it is this complex that is used by archigregarines for the uptake of nutrients by piercing the host cell. ''Gregarina garnhami'' stays present as a extracellular organism in the host, not penetrating the cell membrane of the host. The effect of ''G. garnhami'' on the host is not terribly severe but it is one of the few gregarines which are pathogenic. Harry 1970 found it to cause weight loss but not
molting In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is the manner in which an animal routinely casts off a part of its body (often, but not always, an outer ...
or feeding inhibition. ''Gregarina garnhami'' is used for
biocontrol Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also invo ...
of ''
Schistocerca gregaria The desert locust (''Schistocerca gregaria'') is a species of locust, a periodically swarming, short-horned grasshopper in the family Acrididae. They are found primarily in the deserts and dry areas of northern and eastern Africa, Arabia, and sou ...
'' - adults and
nymph A nymph ( grc, νύμφη, nýmphē, el, script=Latn, nímfi, label=Modern Greek; , ) in ancient Greek folklore is a minor female nature deity. Different from Greek goddesses, nymphs are generally regarded as personifications of nature, are ty ...
s - and may be appropriate for deployment in
Nepal Nepal (; ne, नेपाल ), formerly the Federal Democratic Republic of Nepal ( ne, सङ्घीय लोकतान्त्रिक गणतन्त्र नेपाल ), is a landlocked country in South Asia. It is mai ...
.


References


Further reading

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External links


Tree of Life Gregarina

Gregarina Movies
{{Taxonbar, from=Q89355358 Conoidasida Endoparasites