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[08/25/2004]
Human to Anopheles transmission |
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 The transmission
from human to the Anopheles mosquito and the development of malaria
in the vector
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1. Life-cycle of the parasite in the vector of transmission:
The infection of a female Anopheles mosquito starts with a blood
meal taken from a human carrier of malaria-infected gametocytes. If
parasites at other stages of their development are ingested by the
mosquito, they are killed and digested. The gametocyte stage is
haploid, as are all the stages in Man. It is present in the
peripheral blood in both male and female forms, ordinarily in the
proportion of one male for three females.
The gametocytes starts a process of activation inside the stomach
of the mosquito. They take on a spherical shape and the
erythrocytic plasmatic membrane ruptures. The activated female
gametocyte is called a macrogamete. The male gametocyte undergoes
an exflagellation, a process during which flagella grow out from
the original cell (up to eight, but ordinarily from four to six),
which subsequently break away from the body of the activated male
gametocyte and constitute as many microgametes. All these
transformations occur ten or so minutes after the blood meal. The
male gamete moves under its own power, meets a female gamete which
it subsequently penetrates. This fecundation process ends with the
formation of a diploid egg of spherical shape. A dozen or so hours
after the infesting meal, this egg then becomes an ookinete, via an
intermediate stage referred to as a "retort": during which what is
not yet an ookinete grows from what is already no longer an egg.
This ookinete is elongated in shape and presents the
characteristics of an invasive stage. It moves under its own power
towards the edge of the stomach, penetrates through the peritrophic
envelope (extracellular structure produced by the stomach and
enveloping the whole of the food vacuole), then forces its way
through the epithelial cells to the outer surface of the stomach to
finally come to a halt under the vitreous lamella of the
epithelium. There, the ookinete rounds up to a small sphere called
an oocyst. This oocyst grows considerably for around ten days
undergoing numerous nuclear divisions, the first of which is
reductional. On the inside of the oocyst some
10,000 sporozoïtes develop and when mature are released, when the
oocyst ruptures, to find themselves in the hemolymph of the
mosquito, then throughout its body cavity. These trophozoïtes have
a positive tropism for the salivary glands into which they
penetrate, passing through the epithelium of the salivary glands
and ending up in the salivary duct, ready to be injected with the
saliva during the next blood meal. It would seem to be accepted
that an Anopheles mosquito with sporozoites in its salivary glands
remains infective throughout its life.
| 2. The factors involved in the success of the passage and the development of the parasite in the vector are numerous and their interaction complex.
At least six factors can be individualized: · The temperature is
to be found at the top of the list of these factors: below 18°C and
above 35°C, the sporogonic development
of Plasmodium falciparum is halted.
At temperatures from 20°C, 24°C and 30°C, it is respectively from
20, 11 and 9 days. · The
gametocytic density is also
a factor with an obvious and positive
influence on the infection of the vectors. · The state of immunity of the human carrier
of gametocytes: antibodies directed against the parasite
at the precocious stages of its sporogonic development may block,
partially or even totally, the transmission of the parasite towards
the mosquito, inside the latter. · The drepanocytic status of the gametocyte
carrier: hemoglobin AS favors the passage of the parasite
to the mosquito. · The age of the gametocyte: an age
of 3 to 5 days, after the observation of the first exflagellations,
would seem optimum. · The sex ratio of the gametocytes:
a relatively high proportion of males is favorable to the infection
of the mosquitoes. |
| 3. Is the parasite pathogenic for the mosquito?
In the current state of knowledge, only a negative response can
be given to this question, at least for Anopheles
gambiae, which would seem to tolerate
perfectly Plasmodium falciparum. The
sporogonic development of the parasite and its continued presence
at the sporozoïte stage in the salivary glands would not seem to
have any repercussions on the survival of the infected mosquito. A
low teratogenic effect should not, however, be excluded, but would
be without any major epidemiological implication. Such an effect
would explain, for example, why in Nature only rarely is an
Anopheles mosquito observed with a large number of oocysts, while
in the insectarium, this phenomenon is observed frequently.
Moreover, there is the assurance that certain plasmodial species
which do not parasitize humans are effectively pathogenic for their
vectors.
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