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n\paperw12240\paperh15840\pard\sb0\sl-240{\bkmkstart Pg1}{\bkmkend Pg1}\par\pard
\ql \li10320\sb0\sl-276\slmult0 \par\pard\ql\li10320\sb195\sl-276\slmult0 \up0 \
expndtw-3\charscalex100 \ul0\nosupersub\cf1\f2\fs24 1 \par\pard\li1800\sb0\sl-23
0\slmult0\par\pard\li1800\sb216\sl-230\slmult0\fi0\tx6119\tx6908 \up0 \expndtw-3
\charscalex100 \ul0\nosupersub\cf2\f3\fs20 2004 R. Susanti\tab \up0 \expndtw-3\c
harscalex100 Posted\tab \up0 \expndtw-3\charscalex100 27 November 2004\par\pard\
ql \li1800\sb1\sl-213\slmult0 \up0 \expndtw-3\charscalex100 Makalah Individu \pa
r\pard\ql \li1800\ri7050\sb17\sl-226\slmult0 \up0 \expndtw-3\charscalex100 Penga
ntar Falsafah Sains (PPS702) \line \up0 \expndtw-3\charscalex100 Program Pasca S
arjana/S3 \line \up0 \expndtw-3\charscalex100 Institut Pertanian Bogor \line \up
0 \expndtw-3\charscalex100 November 2004 \par\pard\ql \li1800\sb0\sl-230\slmult0
\par\pard\ql\li1800\sb1\sl-230\slmult0 \up0 \expndtw-2\charscalex100 Dosen: \pa
r\pard\qj \li1800\ri5279\sb19\sl-220\slmult0 \up0 \expndtw-3\charscalex100 Prof.
Dr. Ir. Rudy C. Tarumingkeng (Penanggung jawab) \up0 \expndtw-3\charscalex100 P
rof. Dr. Ir. Zahrial Coto \par\pard\ql \li1800\sb12\sl-230\slmult0 \up0 \expndtw
-3\charscalex100 Dr. Hardjanto, MS \par\pard\ql \li2134\ri1870\sb290\sl-400\slmu
lt0\tx2980 \up0 \expndtw0\charscalex97 \ul0\nosupersub\cf4\f5\fs32 DEFEK ERITROS
IT SEBAGAI FAKTOR RESISTENSI \line\tab \up0 \expndtw0\charscalex98 ALAMI TERHADA
P PENYAKIT MALARIA \par\pard\ql \li5829\sb0\sl-253\slmult0 \par\pard\ql\li5829\s
b69\sl-253\slmult0 \up0 \expndtw-3\charscalex100 \ul0\nosupersub\cf3\f4\fs22 Ole
h: \par\pard\ql \li5414\sb0\sl-322\slmult0 \par\pard\ql\li5414\sb48\sl-322\slmul
t0 \up0 \expndtw-3\charscalex100 \ul0\nosupersub\cf5\f6\fs28 R. Susanti \par\par
d\ql \li5163\sb75\sl-253\slmult0 \up0 \expndtw-3\charscalex100 \ul0\nosupersub\c
f3\f4\fs22 B063040041/SVT \par\pard\ql \li4836\sb38\sl-264\slmult0 \up0 \expndtw
-3\charscalex100 susan_veter@yahoo.com \par\pard\ql \li1800\sb0\sl-253\slmult0 \
par\pard\ql\li1800\sb113\sl-253\slmult0 \up0 \expndtw-3\charscalex100 Latar Bela
kang \par\pard\qj \li1800\ri1624\sb0\sl-310\slmult0\fi720 \up0 \expndtw0\charsca
lex103 \ul0\nosupersub\cf6\f7\fs22 Malaria adalah penyakit parasiter yang sampai
saat ini belum dapat diatasi \line \up0 \expndtw0\charscalex100 secara sempurna
. Berdasarkan laporan WHO (2000), terdapat lebih dari 2400 juta \line \up0 \expn
dtw0\charscalex100 penduduk atau \up0 \expndtw0\charscalex102 40% dari pendud
uk dunia tinggal di daerah endemis malaria. \par\pard\qj \li1799\ri1625\sb

0\sl-305\slmult0\fi0 \up0 \expndtw0\charscalex100 Prevalensi penyakit malaria di

seluruh dunia diperkirakan lebih dari 250 juta setiap \line \up0 \expndtw0\char
scalex103 tahunnya, dengan angka kematian mencapai 2 juta penduduk per tahun. Da
ri 300-\line \up0 \expndtw-1\charscalex100 500 kasus malaria, paling banyak dise
babkan oleh Plasmodium falciparum (Warrel et \line \up0 \expndtw-2\charscalex100
al., \up0 \expndtw0\charscalex106 1990). Di Indonesia, sampai saat ini angka ke
sakitan penyakit malaria masih \line \up0 \expndtw-1\charscalex100 cukup tinggi,
terutama di daerah luar Jawa dan Bali. Namun, kini di daerah Jawa dan \line \up
0 \expndtw0\charscalex100 Bali juga terjadi peningkatan jumlah penderita malaria
, karena banyaknya pengungsi \line \up0 \expndtw0\charscalex100 yang berasal d
ari daerah yang dilanda konflik. Pengungsian tersebut berperan \line \up0
\expndtw0\charscalex106 sebagai media penyebaran malaria dari daerah endemis ke
daerah non-endemis \line \up0 \expndtw-3\charscalex100 (Fahmi, 2001). \par\pard
\qj \li1799\ri1624\sb10\sl-305\slmult0\fi720 \up0 \expndtw0\charscalex106 Malari
a disebabkan oleh parasit golongan protozoa, yaitu Plasmodium sp. \line \up0 \ex
pndtw0\charscalex100 Protozoa ini mempunyai hospes definitif nyamuk Anopheles sp
. Infeksi Plasmodium \line \up0 \expndtw-1\charscalex100 sp pada manusia dimulai
ketika nyamuk Anopheles sp pembawa sporozoit Plasmodium \line \up0 \expndtw0\ch
arscalex105 sp menggigit manusia. Sporozoit yang masuk melalui gigitan nyamuk, s
elanjutnya \line \up0 \expndtw0\charscalex103 melalui sistem sirkulasi menuju ha
ti dan memasuki stadium intrasel hati. Di dalam \line \up0 \expndtw0\charscalex1
03 hepatosit, sporozoit mengalami replikasi aseksual membentuk merozoit. Merozoi
t \line \up0 \expndtw-1\charscalex100 selanjutnya menyerbu dan masuk ke sel-sel
eritrosit, dan dimulailah periode intrasel \line \up0 \expndtw-3\charscalex100 e
ritrosit \up0 \expndtw0\charscalex104 (Wakelin, 1988; Wiser, 2004). Di dalam eri
trosit, merozoit melakukan \line \up0 \expndtw0\charscalex102 metabolisme akti
f termasuk mengingesti sitoplasma eritrosit hospes dan \par\pard\sec
t\sectd\fs24\paperw12240\paperh15840\pard\sb0\sl-240{\bkmkstart Pg2}{\bkmkend Pg
2}\par\pard\ql \li10320\sb0\sl-276\slmult0 \par\pard\ql\li10320\sb195\sl-276\slm
ult0 \up0 \expndtw-3\charscalex100 \ul0\nosupersub\cf1\f2\fs24 2 \par\pard\qj \l
i1800\sb0\sl-300\slmult0 \par\pard\qj\li1800\ri1625\sb145\sl-300\slmult0 \up0 \e
xpndtw0\charscalex104 \ul0\nosupersub\cf6\f7\fs22 menghancurkan hemoglobin menja
di asam amino. Akibatnya, penderita mengalami \up0 \expndtw-3\charscalex100 geja
la anemia (Hyde, 1990). \par\pard\ql \li2520\sb39\sl-253\slmult0\tx4994 \up0 \ex
pndtw-2\charscalex100 Stadium intraseluler \tab \up0 \expndtw0\charscalex100 (
dalam sel hepatosit dan eritrosit) merupakan \par\pard\qj \li1800\ri16
25\sb24\sl-306\slmult0 \up0 \expndtw-1\charscalex100 mekanisme parasit untuk men
ghindar dari sistem imunitas hospes. Dengan berada di \up0 \expndtw0\charscalex1
00 dalam sel hospes, komponen-komponen pertahanan tubuh hospes tidak mengenalnya
\up0 \expndtw0\charscalex104 sebagai bahan asing. Stadium intrasel eritrosit, m
erupakan stadium paling toksik \up0 \expndtw0\charscalex105 dari merozoit bagi h
ospes. Stadium ini diawali dengan proses invasi merozoit ke \up0 \expndtw0\chars
calex107 dalam eritrosit, diikuti pertumbuhan dan replikasi merozoit di dalam er
itrosit. \up0 \expndtw0\charscalex102 Keberhasilan mekanisme invasi dan pertumbu
han merozoit sangat ditentukan oleh \up0 \expndtw-3\charscalex100 kondisi eritro
sit sebagai habitatnya. \par\pard\qj \li1800\ri1624\sb0\sl-306\slmult0\fi720 \up
0 \expndtw-2\charscalex100 Beberapa defek eritrosit secara herediter disebutkan
merupakan pertahanan \line \up0 \expndtw0\charscalex100 alami terhadap malaria.
Defek pada komponen membran eritrosit secara herediter, \line \up0 \expndtw0\cha
rscalex102 seperti defisiensi glikoporin dan antigen duffy berturut-turut dapat
menghambat \line \up0 \expndtw0\charscalex106 invasi P. falciparum dan P. vivax.
Defisiensi enzim G6PD, dan mutasi hemoglobin \line \up0 \expndtw0\charscalex100
seperti talasemia, HbS, dan HbC akan mengganggu pertumbuhan dan replik
asi \line \up0 \expndtw0\charscalex105 merozoit. Bagaimana hal ini dapat terjadi
? Tulisan berikut ini akan menguraikan \line \up0 \expndtw0\charscalex102 mekani
sme resistensi alami dari defek eritrosit terhadap malaria, sehingga dapat \line
\up0 \expndtw0\charscalex100 dipahami bahwa pada penderita defek eritrosit
justru tahan terhadap infeksi \line \up0 \expndtw0\charscalex103 malaria. U
ntuk memudahkan memahami mekanisme tersebut, berturut-turut akan \line \up0 \exp
ndtw-2\charscalex100 diuraikan siklus hidup Plasmodium sp. dan metabolisme eritr

osit normal. \par\pard\ql \li1800\sb0\sl-253\slmult0 \par\pard\ql\li1800\sb99\sl

-253\slmult0 \up0 \expndtw-3\charscalex100 \ul0\nosupersub\cf3\f4\fs22 Siklus Hi
dup Plasmodium sp \par\pard\qj \li1799\ri1625\sb5\sl-305\slmult0\fi720 \up0 \exp
ndtw-1\charscalex100 \ul0\nosupersub\cf6\f7\fs22 Siklus hidup Plasmodium sp meli
batkan insekta sebagai hospes definitiv (yaitu \line \up0 \expndtw0\charscalex10
5 nyamuk) dan hospes intermedier yaitu vertebrata (termasuk manusia) (Wakelin, \
line \up0 \expndtw-1\charscalex100 1988). Dalam hospes definitiv, plasmodium men
galami stadium seksual dan sporogoni, \line \up0 \expndtw0\charscalex100 sedangk
an dalam hospes intermedier mengalami stadium aseksual yaitu intrasel hati \line
\up0 \expndtw-1\charscalex100 dan eritrosit. Dalam siklus hidupnya, Plasmodium
sp melalui 3 stadium aseksual, yaitu \line \up0 \expndtw0\charscalex100 eksoerit
rositik skizogoni, stadium eritrositik skizogoni dan sporogoni, menghasilkan \li
ne \up0 \expndtw0\charscalex105 bentuk invasif merozoit dan sporozoit. Seperti p
ada spesies apikompleksa pada \line \up0 \expndtw0\charscalex100 umumnya, semua
stadium invasif Plasmodium sp mempunyai karakteristik pada \line \up0 \e
xpndtw0\charscalex103 organela apeknya (Wiser, 2004). Secara keseluruhan, siklu
s hidup Plasmodium sp \line \up0 \expndtw-1\charscalex100 terdiri dari \up0 \ex
pndtw0\charscalex100 4 stadium yaitu stadium intrasel hati, stadium intra
sel eritrosit, \line \up0 \expndtw0\charscalex104 stadium seksual dan sporogoni
(Wiser, 2004). Stadium intrasel merupakan salah \line \up0 \expndtw-2\charscale
x100 satu strategi parasit ini menghindar dari sistem pertahanan tubuh hospes. \
par\pard\qj \li1799\ri1625\sb15\sl-305\slmult0\fi720 \up0 \expndtw0\charscalex10
5 Infeksi Plasmodium sp pada manusia dimulai ketika nyamuk Anopheles sp \line \u
p0 \expndtw0\charscalex100 pembawa sporozoit Plasmodium sp menggigit manusia. Sp
orozoit yang masuk melalui \line \up0 \expndtw0\charscalex106 gigitan nyamuk, se
lanjutnya melalui sistem sirkulasi menuju hati dan memasuki \line \up0 \expndtw
0\charscalex103 stadium intrasel hati (sporozoit masuk ke sel-sel hepatosit hati
) (Wakelin, 1988). \line \up0 \expndtw0\charscalex100 Pemasukan sporozoit ke
dalam hepatosit diperantarai oleh ikatan antara \line \up0 \expndtw
0\charscalex104 circumsporozoit protein (CSP) pada membran sporozoit dengan prot
ein membran \line \up0 \expndtw0\charscalex102 hepatosit yaitu heparin sulfat pr
oteoglikan (Sinnis and Sim, 1997). Selama berada \line \up0 \expndtw0\charscalex
100 dalam hepatosit (stadium intrasel), sporozoit mengalami skizogoni (replikasi
secara \par\pard\sect\sectd\fs24\paperw12240\paperh15840\pard\sb0\sl-240{\bkmks
tart Pg3}{\bkmkend Pg3}\par\pard\ql \li10320\sb0\sl-276\slmult0 \par\pard\ql\li1
0320\sb195\sl-276\slmult0 \up0 \expndtw-3\charscalex100 \ul0\nosupersub\cf1\f2\f
s24 3 \par\pard\qj \li1800\sb0\sl-300\slmult0 \par\pard\qj\li1800\ri1626\sb145\s
l-300\slmult0 \up0 \expndtw-1\charscalex100 \ul0\nosupersub\cf6\f7\fs22 aseksual
), yaitu pembelahan inti sporozoit tanpa diikuti sitogenesis. Proses skizogoni \
line \up0 \expndtw-1\charscalex100 menghasilkan schizont \up0 \expndtw-1\chars
calex100 (disebut schizont eksoeritrositik/preeritrositik) yang \par\pard\
qj \li1799\ri1626\sb0\sl-310\slmult0\fi0 \up0 \expndtw-1\charscalex100 mengandun
g ribuan merozoit. Merozoit-merozoit dalam schizont eksoeritrositik \line
\up0 \expndtw0\charscalex103 selanjutnya keluar dari hepatosit menuju sistem sir
kulasi (Wakelin, 1988; Wiser, \line \up0 \expndtw-1\charscalex100 2004). Selama
stadium intrasel hati, P. vivax dan P. ovale mengalami stadium dorman \line \up0
\expndtw0\charscalex104 dengan membentuk hipnozoit. Hipnozoit tersebut akan rea
ktif setelah beberapa \line \up0 \expndtw-2\charscalex100 minggu, bulan atau tah
un dan akan mengalami replikasi aseksual (Wiser, 2004). \par\pard\qj \li1799\ri1
624\sb0\sl-306\slmult0\fi720 \up0 \expndtw-1\charscalex100 Merozoit selanjutnya
menyerbu dan masuk ke sel-sel eritrosit, dan dimulailah \line \up0 \expndtw-1\ch
arscalex100 periode intrasel eritrosit (Wakelin, 1988). Stadium intraseluler eri
trosit melibatkan \line \up0 \expndtw0\charscalex100 interaksi seluler antara er
itrosit-merozoit. Periode ini disebut juga periode tropik \line \up0 \expndtw0\c
harscalex104 (Wiser, 2004), kemungkinan karena pada stadium ini menyebabkan geja
la demam \line \up0 \expndtw0\charscalex106 dengan suhu tubuh mencapai 41\ul0\su
per\cf7\f8\fs21 o\ul0\nosupersub\cf6\f7\fs22 C pada penderitanya. Merozoit dalam
eritrosit \line \up0 \expndtw0\charscalex104 mengalami pertumbuhan dan pembelah
an. Pada awalnya merozoit dalam eritrosit \line \up0 \expndtw0\charscalex102 ber
bentuk cincin \up0 \expndtw0\charscalex106 (ring form) dan diikuti dengan pertam

bahan volume merozoit \par\pard\qj \li1799\ri1625\sb0\sl-306\slmult0 \up0 \expnd

tw-1\charscalex100 (membentuk tropozoit) (Wakelin, 1988; Wiser, 2004). Pertambah
an volume merozoit \up0 \expndtw0\charscalex102 terjadi karena merozoit mela
kukan metabolisme aktif termasuk mengingesti \up0 \expndtw0\charscalex104 s
itoplasma eritrosit hospes dan menghancurkan hemoglobin menjadi asam amino. \up0
\expndtw-3\charscalex100 Akibatnya, penderita mengalami gejala anemia (Hyde, 19
90). \par\pard\ql \li2519\sb38\sl-253\slmult0\tx9589 \up0 \expndtw-1\charscalex1
00 Selanjutnya tropozoit melakukan skizogoni membentuk schizont \tab \up0 \
expndtw-2\charscalex100 (disebut \par\pard\qj \li1799\ri1625\sb5\sl-305\slmult0
\up0 \expndtw0\charscalex102 schizont eritrositik), yang kurang lebih mengandung
32 merozoit (Wakelin, 1988). \line \up0 \expndtw0\charscalex100 Merozoit-merozo
it dalam schizont selanjutnya dilepaskan dari eritrosit dengan \line \up0
\expndtw0\charscalex100 merusak dinding eritrosit, dan menginfeksi eritrosit la
in. Dalam eritrosit, merozoit \line \up0 \expndtw0\charscalex105 ini akan mengal
ami pertumbuhan dan perkembangan yang sama seperti merozoit \line \up0 \expndtw0
\charscalex104 sebelumnya (Wakelin, 1988; Wiser, 2004). Demam tinggi pada pender
ita malaria \line \up0 \expndtw0\charscalex103 dikarenakan multiplikasi mero
zoit, rupturnya eritrosit ketika melepaskan \line \up0 \expndtw-1\charsc
alex100 merozoitnya dan re-invasi ke eritrosit lain (Hyde, 1990). Ketika pelepas
an merozoit, \line \up0 \expndtw0\charscalex102 pigmen dan semua produk meta
bolitnya juga dilepaskan ke sirkulasi sehingga \line \up0 \expndtw-3\charsc
alex100 menginduksi perubahan-perubahan patologis hospes (Wakelin, 1988). \par\p
ard\qj \li1799\ri1624\sb15\sl-306\slmult0\fi720 \up0 \expndtw0\charscalex102 Pad
a infeksi P. falciparum, eritrosit yang mengandung merozoit, selain \up0
\expndtw0\charscalex104 ditemukan di darah perifer juga ditemukan menempel pada
sel endotel pembuluh \up0 \expndtw0\charscalex105 darah kapiler organ-organ vis
ceral. Agregasi dan blokade merozoit pada kapiler \up0 \expndtw0\charscalex100 t
ersebut menyebabkan kerusakan ginjal, paru dan otak sehingga menyebabkan
\up0 \expndtw0\charscalex103 kematian (Wakelin, 1988; Hyde, 1990). Penempelan m
erozoit pada endotel kapiler \up0 \expndtw0\charscalex106 darah merupakan salah
satu mekanisme parasit untuk bersembunyi dari sistem \up0 \expndtw-3\charscalex1
00 pertahanan tubuh hospes (Wiser, 2004). \par\pard\qj \li1799\ri1625\sb0\sl-306
\slmult0\fi720 \up0 \expndtw0\charscalex106 Selain replikasi secara aseksual, pa
rasit juga mengalami stadium seksual \line \up0 \expndtw-1\charscalex100 dengan
terlebih dahulu membentuk makrogametosit dan mikrogametosit. Stadium ini \line \
up0 \expndtw0\charscalex102 disebut preseksual (Hyde, 1990). Gametosit (mikrogam
et dan makrogamet) adalah \line \up0 \expndtw0\charscalex105 merozoit yang besar
dan hampir memenuhi eritrosit, dan setiap eritrosit hanya \line \up0 \expndtw0\
charscalex108 mengandung 1 merozoit (Wiser, 2004). Pada saat nyamuk Anopheles sp
betina \line \up0 \expndtw0\charscalex103 menggigit manusia yang eritrositnya m
engandung gametosit, gametosit teringesti \line \up0 \expndtw0\charscalex100 ole
h nyamuk, menginduksi gametogenesis dan keluar dari eritrosit. Beberapa faktor \
par\pard\sect\sectd\fs24\paperw12240\paperh15840{\shp{\*\shpinst\shpleft3240\shp
top4811\shpright8462\shpbottom10008\shpbxpage\shpbypage\shpwr3\shpfblwtxt1\shpz9
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f3ae7ff6f0f86ffdbdff00083fc72f07ff00c4c3c39ac68ffd83e21d6341ec7fe69bfbfafd3ebd3f
0ac42faae778bc35ff00e4558cf277ff003f9ee757fcc93fedefd4f48f8b5e09d3741fdb7fe0878c
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4bfb634ff11f8c358f156bde1ed63d0fe19ffeb7bd7d01e0ab3f07fed69fb3af85ff00e130fed8ff
00848f47ff00890ff6c7fcccfa1f8d7e1f7e63af5e95f23f8f35efb67c1fd3fe39683e20d1fc2ff1
fbe1feb1ff00081fc50d634b1ff21ce71e9cd79b88ff00a095ff00315fed9eb6fc0f473076ff0085
adaddaddbeff00f807eac7fc1337c6d0ebdfb28f81f4cfb459ea1ff08bf8c3c55a0eae3fe807ff00
33e7fede63f5a83c53fb17f8e3e3af897c613fed63fb44788fe347c10d43c61fdbbf0fff00648f07
781c7c17f814be173fe83e05f0e7c77fb27c42bef8a9fb47ba82c7e2efc38f8a7f14c7ecf9f17dc0
23f6714601c7937fc11ff48f1559fc26f89177a91b3ff843f50f1e67c2e3fe9f0e7fe13c3c60f247
8407e1ed5fb11077fc7fa57eeb93ff00b565d85b74feafd3faf43c4a1ff22e7f3fc91c6f86fc2be1
af0768561e1bf0ae81a4787bc2be19d27fb0747d1b4cd10f867c31a1f86768185049006172554007
e66232493e65fb60ff00c9adfc79ff00b271e2cffd379afa1ebe78fdb07fe4d6fe3cff00d938f167
fe9bcd74e27fddf15ff607fe407e437fc13df47fb1fc1ff146a5ff00417f1e7a1fc3a7f91f99afba
2be27fd80bfe4829ff00b1f3c57ffa0d7db15fcf188ff919e1bfae8cfb6e18ff009177ddf9851451
5ce7d505145140051451400514514005145140051451ff00838a002bf1abfe0e02ff0094437ed71f
4f80bffad5ff00b3c57ecaff00c4b7fea31fa57e3cff00c1c1dff2887fdb23fddfd9d7ff005abff6
79af4b27ff00919e53ff0061d803e7f36ff9177157fd861fb0ff00f04b3ff9463ffc139bfecc43f6
43ff00d67ef87b451ff04b3ff9463ffc139bfecc43f643ff00d67ef87b457f42ad97a2fc91f903dd
fabfcd9fccaffc1ab3fb61fec9bfb34ffc13e7e2df817e3efed3ff00b39fc09f17f88ff6c7f8a1e2
9f0e683f19be2ff803e176b1a87864fc0efd9e2c1bc5da78f1e6a001449ac26854a82098980e54d7
f5063fe0a9ff00f04be3d3fe0a3dfb089fa7ed87f000ff00ef43aff174bcebf80fe62a95b7defcff
0090af32129ca0e7ced59ec92fcd9d786ad528d0c47b3938f3bd7ad9ded757f4fd0ff698ff0087aa
7fc1327fe922ff00b07ffe25afc0ff00fe78547fc3d53fe0993ff4917fd83fff0012d7e07fff003c
2aff00175a2b4fed1a9ff3ea8ffe03ff0004e7fed0c67fcfeffc963fe67fb457fc3d53fe0993ff00
4917fd83ff00f12d7e07ff00f3c2a3fe1ea9ff0004c9ff00a48bfec1ff00f896bf03ff00f9e157f8
bad147f68d4ff9f547ff0001ff00821fda18cff9fdff0092c7fccff68aff0087aa7fc1327fe922ff
00b07ffe25afc0ff00fe78547fc3d53fe0993ff4917fd83fff0012d7e07fff003c2aff00175a28fe
d1a9ff003ea8ff00e03ff043fb4319ff003fbff258ff0099fed15ff0f54ff8264ffd245ff60fff00
c4b5f81fff00cf0a8ff87aa7fc1327fe922ffb07ff00e25afc0fff00e7855fe2eb451fda353fe7d5
1ffc07fe087f68633fe7f7fe4b1ff33fd74bf6b5fdacbfe094bfb517c3e1a0cdff000516fd84347f
1cf860ff006ef817c5ff00f0d4ff00043fe296bece028ff8b89c9e78039fe55f91fe0efdbfff0065
4f841e01f895f00fe2d7ed3bfb38fc50f09eada405f046b5f0bff68af821e28018e70a4ffc2d5c02
70700e09c76db5fe746ff74fe1fcc5581f79bfe03fcabe6f89387f039e548d6c54abd3ad2c2a87b4
a15142d7bd9d9a77b5b5d7d2c690cdb30a69b5886ecbac52d975b3daff00858ff454f827fb537847
e135b7d9bc07fb76ff00c13e74fd1bc62dfdb9abf873c65fb68fecc1ff0008de86d9c6ddbff0b573
bb760000641e00cd7a07d9ff0062ff00ec7d634dbbff0082bb7fc1350ffc253acf85b5dd7c8fda67
e10726c70580ff008babdb2091838c826bfcd568ae6a7c2b96c7f89571b88ffafd88e6ff00db3fe0
791d1f5bc47fcfc7f77fc13fd847f66ffdbebfe09abf04be06f81be15eb1ff00052ffd84358bef08
733ea1a67ed6bf043907c402f7fe8a2608239cf4e41cd7d03ff0f54ff8261671ff000f1ffd8433e9
ff000d89f0473ffab12bfc5acfde5ff817f2a6ff00cb4ff3fddafb64bead430aa9696c1b57eb64ed
e97662aa54693e67aa5dbb1fed2dff000f54ff008261ff00d247ff00611ffc4c4f823ffcf12be7ff
00da57fe0a69ff0004ddf107ecf7f1c341f0ff00edfbfb14eb5ae6b1f0b7c6d65a3e8fa07ed4df04
3c51e27d6efdfc1cdb56cc8f8867e66da76af56c1c03835fe3d69f747e3fccd03ef37fc07f956552
b54a94b110726aed2befbb6b6f2b750752a24df33d2e7fa837ec4ffb7b7ec35e13f8003c3be31fdb
4ff64ef0b6a87c79e29d7c693aafed2ff047c2ea770f87db793f1531f379726dc7ded8f8e5588fab
7fe1e3dff04ecffa3eff00d8cbff00129fe07fff003d6aff002587fbc7f0fe429b5f0d3e0dcaea51
a31f6f8f8a85ecd622377676d7dcd3e5e5d8f4f2de25cdb2e4beab88e4bbb6aafe5d2c7fad4ffc3c
97fe09dbff0047effb1a7fe254fc10ff00e7a947fc3c97fe09dbff0047effb1a7fe254fc10ff00e7
a95fe4cd4571ff00a9395ffd05663ff8330bff00ccc7b5feb5e3bfe80f2cff00c24ffedcff00599f
f8792ffc13b7fe8fdff634ff00c4a9f821ff00cf528ff8792ffc13b7fe8fdff634ff00c4a9f821ff
00cf52bfc99a8a3fd49caffe82b31ffc1985ff00e660ff005af1dff407967fe127ff006e7faccffc

3c97fe09dbff0047effb1a7fe254fc10ff00e7a947fc3c97fe09dbff0047effb1a7fe254fc10ff00
e7a95fe4cd451fea4e57ff0041598ffe0cc2ff00f3307fad78effa03cb3ff093ff00b73fd667fe1e
4bff0004edff00a3f7fd8d3ff12a7e087ff3d4a3fe1e4bff0004edff00a3f7fd8d3ff12a7e087ff3
d4aff266a28ff5272bff00a0acc7ff0006617ff9983fd6bc77fd01e59ff849ff00db9feb33ff000f
25ff008276ff00d1fbfec69ff8953f043ff9ea51ff000f25ff008276ff00d1fbfec69ff8953f043f
f9ea57f9335147fa9395ff00d05663ff008330bffccc1feb5e3bfe80f2cffc24ff00edcff599ff00
8792ff00c13b7fe8fdff00634ffc4a9f821ffcf528ff008792ff00c13b7fe8fdff00634ffc4a9f82
1ffcf52bfc99a8a3fd49caff00e82b31ff00c1985ffe660ff5af1dff00407967fe127ff6e7facd7f
c3c7bfe09d901c1fdbbff634c8edff000d4ff043d3dfe2afbfd49f5afca4ff0082e0fed99fb1bfc6
9ff82597ed39f0f3e0efed69fb33fc53f1bf8a1be0edde8fe10f85df1fbe1ff8cfc69ae2e9df1bfe
1e5fea04fc3bf8720aa88e35779247f95115a4930aac47f9d95c7de7ff0080ff0025ad2d3ffe3ddb
f0ff00d0debd38f0d6030f2a389a53c446bd3c4616aa9f3c6cdd27a45ab6d2b3bb5aebea7939b711
66b89c830983ab896e8aa895946cdda4bcdef6e8bccff65dff008259ff00ca31ff00e09cdff6621f
b21ffeb3f7c3da28ff008259ff00ca31ff00e09cdff6621fb21ffeb3f7c3da2bef56cbd17e48f9f7
bbf57f9b3fffd9}}}}}
\pard\sb0\sl-240{\bkmkstart Pg4}{\bkmkend Pg4}\par\pard\ql \li10320\sb0\sl-276\s
lmult0 \par\pard\ql\li10320\sb195\sl-276\slmult0 \up0 \expndtw-3\charscalex100 \
ul0\nosupersub\cf1\f2\fs24 4 \par\pard\qj \li1799\sb0\sl-305\slmult0 \par\pard\q
j\li1799\ri1625\sb136\sl-305\slmult0\fi0 \up0 \expndtw0\charscalex100 \ul0\nosup
ersub\cf6\f7\fs22 yang menginduksi gametogenesis adalah turunnya suhu, peningkat
an karbondioksida \up0 \expndtw-1\charscalex100 dan metabolit nyamuk. Mikr
ogamet dan makrogamet selanjutnya mengalami \up0 \expndtw0\charscalex100
fertilisasi di dalam saluran pencernaan nyamuk membentuk zigot. Zigot selanjutn
ya \up0 \expndtw0\charscalex106 berkembang menjadi bentuk motil (disebut ookinet
) dan masuk ke dinding usus. \up0 \expndtw0\charscalex100 Ookinet berkembang
menjadi ookista. Ookista mengalami stadium aseksual \up0 \expndtw0\ch
arscalex103 (sporogoni) sehingga membentuk sporozoit. Sporozoit selanjutnya
migrasi ke \up0 \expndtw0\charscalex105 glandula saliva. Jika nyamuk ini kemu
dian menggigit manusia, siklus akan kembali \up0 \expndtw0\charscalex103 berulan
g seperti semula (Wakelin, 1988; Wiser, 2004). Secara garis besar, siklus \up0 \
expndtw-3\charscalex100 hidup Plasmodium sp terlihat pada Gambar 1. \par\pard\ql
\li1799\sb0\sl-276\slmult0 \par\pard\ql\li1799\sb0\sl-276\slmult0 \par\pard\ql\
li1799\sb0\sl-276\slmult0 \par\pard\ql\li1799\sb0\sl-276\slmult0 \par\pard\ql\li
1799\sb0\sl-276\slmult0 \par\pard\ql\li1799\sb0\sl-276\slmult0 \par\pard\ql\li17
99\sb0\sl-276\slmult0 \par\pard\ql\li1799\sb0\sl-276\slmult0 \par\pard\ql\li1799
\sb0\sl-276\slmult0 \par\pard\ql\li1799\sb0\sl-276\slmult0 \par\pard\ql\li1799\s
b0\sl-276\slmult0 \par\pard\ql\li1799\sb0\sl-276\slmult0 \par\pard\ql\li1799\sb0
\sl-276\slmult0 \par\pard\ql\li1799\sb0\sl-276\slmult0 \par\pard\ql\li1799\sb0\s
l-276\slmult0 \par\pard\ql\li1799\sb0\sl-276\slmult0 \par\pard\ql\li1799\sb0\sl276\slmult0 \par\pard\ql\li1799\sb0\sl-276\slmult0 \par\pard\ql\li1799\sb0\sl-27
6\slmult0 \par\pard\ql\li1799\sb0\sl-276\slmult0 \par\pard\ql\li1799\sb0\sl-276\
slmult0 \par\pard\ql\li1799\sb0\sl-276\slmult0 \par\pard\ql\li1799\sb67\sl-276\s
lmult0 \up0 \expndtw-2\charscalex100 Gambar 1. Siklus hidup Plasmodium sp. ({\fi
eld{\*\fldinst {HYPERLINK "http://whyfiles.org/016skeeter/malaria2.html" }}{\fld
rslt {\ul0\nosupersub\cf11\f12\fs24\ul http://whyfiles.org/016skeeter/malaria2.h
tml}}}\ul0\nosupersub\cf1\f2\fs24 ) \par\pard\ql \li1799\sb0\sl-253\slmult0 \par
\pard\ql\li1799\sb0\sl-253\slmult0 \par\pard\ql\li1799\sb157\sl-253\slmult0 \up0
\expndtw-3\charscalex100 \ul0\nosupersub\cf3\f4\fs22 Metabolisme eritrosit norm
al \par\pard\qj \li1800\ri1626\sb29\sl-300\slmult0\fi719 \up0 \expndtw-1\charsca
lex100 \ul0\nosupersub\cf6\f7\fs22 Meskipun ada 2 bentuk stadium invasif, yaitu
merozoit dan sporozoit, bentuk \line \up0 \expndtw-2\charscalex100 invasif mero
zoit \up0 \expndtw-1\charscalex100 (terutama intra eritrosit) banyak mendapa
t perhatian karena \par\pard\qj \li1800\ri1625\sb0\sl-310\slmult0 \up0 \expndt
w0\charscalex104 banyak menimbulkan efek patologis pada hospes vertebrata (terma
suk manusia). \line \up0 \expndtw0\charscalex106 Merozoit secara cepat dan spesi
fik menginfeksi eritrosit, melibatkan interaksi \line \up0 \expndtw0\charscalex1
00 reseptor dan ligan (Wiser, 2004). Proses invasi merozoit pada eritrosit terdi
ri dari \par\pard\qj \li1800\ri1625\sb0\sl-306\slmult0 \up0 \expndtw0\charscalex
103 4 tahap yaitu pengikatan merozoit pada membran eritrosit, reorientasi eritro
sit, \line \up0 \expndtw-1\charscalex100 pembentukan junction dan pemasukan para

sit (Gratzer and Dluzewski, 1993). Selama \line \up0 \expndtw0\charscalex107 di

dalam eritrosit, merozoit membutuhkan asam amino untuk dibentuk menjadi \line \u
p0 \expndtw-1\charscalex100 protein. Asam amino diperoleh merozoit dari 3 sumber
yaitu sintesis de novo, import \par\pard\sect\sectd\fs24\paperw12240\paperh1584
0\pard\sb0\sl-240{\bkmkstart Pg5}{\bkmkend Pg5}\par\pard\ql \li10320\sb0\sl-276\
slmult0 \par\pard\ql\li10320\sb195\sl-276\slmult0 \up0 \expndtw-3\charscalex100
\ul0\nosupersub\cf1\f2\fs24 5 \par\pard\qj \li1800\sb0\sl-300\slmult0 \par\pard\
qj\li1800\ri1625\sb145\sl-300\slmult0 \up0 \expndtw0\charscalex102 \ul0\nosupers
ub\cf6\f7\fs22 dari hospes dan degradasi hemoglobih eritrosit. Degradasi hemoglo
bin merupakan \up0 \expndtw-3\charscalex100 sumber asam amino utama bagi merozoi
t. \par\pard\qj \li1800\ri1622\sb0\sl-308\slmult0\fi720 \up0 \expndtw-2\charscal
ex100 Mengingat eritrosit merupakan habitat bagi merozoit selama stadium intrase
l \line \up0 \expndtw0\charscalex100 eritrosit, berikut ini diuraikan tentang me
tabolisme eritrosit secara normal, serta \line \up0 \expndtw-1\charscalex100 hem
oglobin sebagai sumber utama asam amino bagi merozoit. Eritrosit manusia tidak \
line \up0 \expndtw0\charscalex102 mempunyai inti, mitokondria dan organela-organ
ela lainnya. Glukosa yang diperoleh \line \up0 \expndtw0\charscalex105 dari ekst
rasel, merupakan satu-satunya molekul utama untuk dioksidasi menjadi \line \up0
\expndtw0\charscalex102 energi melalui glikolisis \up0 \expndtw0\charscalex106 (
Koolman dan Rohm, 2001). Glukosa masuk ke dalam sel \par\pard\qj \li1799\ri1625\
sb0\sl-308\slmult0\fi0 \up0 \expndtw0\charscalex103 dengan bantuan protein gluko
sa-transporter, selanjutnya melalui proses glikolisis \line \up0 \expndtw0\chars
calex106 diubah menjadi 2 molekul asam laktat dan menghasilkan 2 molekul ATP. Ka
rena \line \up0 \expndtw0\charscalex100 eritrosit tidak memiliki mitokondria, mo
lekul hasil proses glikolisis tidak dioksidasi \line \up0 \expndtw0\charscalex10
2 lebih lanjut melalui siklus krebs maupun fosforilasi oksidatif (Murray et al.,
2000; \line \up0 \expndtw0\charscalex100 Marks et al, 2001). ATP yang dihasilka
n terutama menyediakan energi bagi Na+/K\ul0\super\cf7\f8\fs21 +\ul0\nosupersub\
cf6\f7\fs22 -\line \up0 \expndtw-2\charscalex100 ATP-ase untuk menjaga lingkunga
n ion di dalam sel (Koolman dan Rohm, 2001). \par\pard\qj \li1800\ri1625\sb0\sl310\slmult0\fi720 \up0 \expndtw0\charscalex102 Selain melalui jalur glikolisis,
glukosa juga dimetabolisme melalui jalur jalur \line \up0 \expndtw0\charscalex10
2 penthosaphosphate shunt (PPS)/hexosa monophosphate shunt (HMPS) (Murray et \li
ne \up0 \expndtw-2\charscalex100 al., \up0 \expndtw0\charscalex101 2000; Marks
et al., \up0 \expndtw0\charscalex101 2001). Jalur PPS ini merupakan jalur
penting untuk \par\pard\qj \li1799\ri1625\sb0\sl-308\slmult0\fi0 \up0 \expndt
w0\charscalex104 melindungi eritrosit dari kerusakan oksidatif, dengan menghasil
kan nicotinamide \up0 \expndtw0\charscalex103 adenine dinucleotide phosphate-H (
NADPH). Reactif oxygen species (ROS) yaitu \up0 \expndtw0\charscalex100 oksigen
dan derivatnya, selain dapat mengubah hemoglobin menjadi methemoglobin \up0 \exp
ndtw0\charscalex100 juga bersifat sangat toksik bagi membran eritrosit (Koolman
dan Rohm, 2001). ROS \up0 \expndtw0\charscalex102 akan menyebabkan lisis (hemoli
sis) akibat kerusakan membran. Pembentukan ROS \up0 \expndtw0\charscalex100 dapa
t dicegah jika dalam sel terdapat glutation reduktase, yaitu enzim yan
g \up0 \expndtw-2\charscalex100 mengubah GSSG (glutation teroksidasi) menjadi GS
H (glutation). Regenerasi GSH ini \up0 \expndtw-2\charscalex100 memerlukan NADPH
sebagai koenzim (Koolman dan Rohm, 2001). \par\pard\ql \li2340\sb7\sl-253\slmul
t0\tx6614 \up0 \expndtw-1\charscalex100 Glucose-6-phosphate
dehidrogenase \ta
b \up0 \expndtw0\charscalex102 (G6PD)
merupakan
enzim
yang \par\pard\qj
\li1799\ri1625\sb4\sl-306\slmult0\fi0 \up0 \expndtw0\charscalex102 mengkatalisi
s reaksi pertama jalur PPS, yaitu mengkatalisis glukosa-6 fosfat \line \
up0 \expndtw-1\charscalex100 menjadi 6-fosfoglukonolakton (fosfoglukonat) (Koolm
an dan Rohm, 2001). Defisiensi \line \up0 \expndtw0\charscalex104 G6PD mengaki
batkan reaksi selanjutnya dalam PPS terganggu, sehingga \line \up0 \
expndtw0\charscalex103 pembentukan NADPH juga mengalami hambatan. Gagalnya
detoksifikasi ROS \line \up0 \expndtw0\charscalex102 menjadi molekul non to
ksik akibat defisiensi NADPH, menyebabkan eritrosit \line \up0 \expndtw0\ch
arscalex106 mengalami stress oksidatif sehingga mudah lisis (Marks et al., 2001)
. Beberapa \line \up0 \expndtw0\charscalex102 mutasi G6PD menyebabkan eritrosit
dihancurkan sistem fagosit secara prematur, \line \up0 \expndtw-1\charscalex100

sehingga menimbulkan gejala anemia pada individu penderitanya. Bayi penderita ya

ng \line \up0 \expndtw0\charscalex100 baru lahir akan menunjukkan gejala ke
kuningan pada mata, kulit dan membran \line \up0 \expndtw0\charscalex106 mu
kosa
disebabkan
oleh
tingginya
deposisi
bilirubin
pada
jari
ngan \line \up0 \expndtw-3\charscalex100 ({\field{\*\fldinst {HYPERLINK "http://
www.as.ua.edu/ant/bindon/" }}{\fldrslt {\ul0\nosupersub\cf12\f13\fs22\ul http://
www.as.ua.edu/ant/bindon/}}}\ul0\nosupersub\cf12\f13\fs22\ul ant475/g6pd/img003
.jpg\ul0\nosupersub\cf6\f7\fs22 ). \par\pard\qj \li1800\ri1623\sb15\sl-305\slmul
t0\fi786 \up0 \expndtw0\charscalex104 Hemoglobin (Hb) adalah protein utama dalam
eritrosit, berfungsi sebagai \line \up0 \expndtw0\charscalex100 pembawa oksigen
. Penyebutan hemoglobin didasarkan pada komponen penyusunnya, \line \up0 \expndt
w0\charscalex100 yaitu globin dan heme. Hemoglobin orang dewasa merupakan tetram
er globin terdiri \line \up0 \expndtw0\charscalex102 dari 2 rantai \u945? dan 2
rantai \u946? masing-masing sekitar 16 kDa. Setiap subunit globin \line \up0 \ex
pndtw0\charscalex107 membawa
satu
gugus
heme \up0 \expndtw-2\charscalex
100 (molekul \up0 \expndtw0\charscalex104 anorganik
mengandung
besi) \par\
pard\sect\sectd\fs24\paperw12240\paperh15840\pard\sb0\sl-240{\bkmkstart Pg6}{\bk
mkend Pg6}\par\pard\ql \li10320\sb0\sl-276\slmult0 \par\pard\ql\li10320\sb195\sl
-276\slmult0 \up0 \expndtw-3\charscalex100 \ul0\nosupersub\cf1\f2\fs24 6 \par\pa
rd\qj \li1800\sb0\sl-305\slmult0 \par\pard\qj\li1800\ri1624\sb136\sl-305\slmult0
\up0 \expndtw-1\charscalex100 \ul0\nosupersub\cf6\f7\fs22 ({\field{\*\fldinst {
HYPERLINK "http://www.brainyencyclopedia.com/encyclopedia/h/he/hemoglobin.html./
" }}{\fldrslt {\ul0\nosupersub\cf12\f13\fs22\ul http://www.brainyencyclopedia.co
m/encyclopedia/h/he/hemoglobin.html\ul0\nosupersub\cf6\f7\fs22 .}}}\ul0\nosupers
ub\cf6\f7\fs22 ). Kurang \line \up0 \expndtw-2\charscalex100 lebih 80% asam am
ino globin membentuk heliks alpha yang dinamai dengan huruf A-H \line \up0 \expn
dtw0\charscalex105 (Koolman dan Rohm, 2001). Oksigen berikatan pada atom besi g
ugus heme yang \line \up0 \expndtw0\charscalex100 bervalensi dua dan terletak di
pusat (Koolman dan Rohm, 2001; Murray et al., 2000; \line \up0 \expndtw-3\chars
calex100 Marks et al., 2001). \par\pard\ql \li1799\sb0\sl-253\slmult0 \par\pard\
ql\li1799\sb105\sl-253\slmult0 \up0 \expndtw-2\charscalex100 \ul0\nosupersub\cf3
\f4\fs22 Mekanisme Defek Eritrosit Sebagai Faktor Resistensi Alami terhadap Mala
ria \par\pard\qj \li1799\ri1625\sb4\sl-306\slmult0\fi720 \up0 \expndtw0\charscal
ex103 \ul0\nosupersub\cf6\f7\fs22 Stadium intrasel eritrosit, merupakan stadium
paling toksik dari merozoit \line \up0 \expndtw0\charscalex102 bagi hospes. Stad
ium ini diawali dengan proses invasi merozoit ke dalam eritrosit, \line \up0 \ex
pndtw0\charscalex104 diikuti pertumbuhan dan replikasi merozoit di dalam
eritrosit. Keberhasilan \line \up0 \expndtw-2\charscalex100 mekanisme invasi da
n pertumbuhan merozoit sangat ditentukan oleh kondisi eritrosit \line \up0 \expn
dtw0\charscalex102 sebagai habitatnya. Mekanisme invasi melibatkan interaksi mem
bran eritrosit dan \line \up0 \expndtw0\charscalex100 membran merozoit, sehing
ga defek pada membran eritrosit akan menyebabkan \line \up0 \expndtw0\char
scalex102 berkurangnya kekuatan interaksi dan invasi gagal dilakukan. Defek pada
komponen \line \up0 \expndtw-1\charscalex100 membran eritrosit secara herediter
, seperti defisiensi glikoporin dan antigen duffy \line \up0 \expndtw0\charscale
x103 berturut-turut dapat menghambat invasi P. falciparum dan P. vivax. Pertumbu
han \line \up0 \expndtw0\charscalex100 dan perkembangan merozoit selama di
dalam eritrosit, sangat ditentukan oleh \line \up0 \expndtw0\charscalex100 k
ondisi sitoplasma eritrosit. Jika di dalam sitoplasma eritrosit mengandung fakto
r-\line \up0 \expndtw0\charscalex102 faktor yang memberi suasana tidak nyam
an bagi merozoit, pertumbuhan dan \line \up0 \expndtw0\charscalex100 perkemb
angannya terganggu bahkan terhambat. Defisiensi enzim G6PD, dan mutasi \line \up
0 \expndtw0\charscalex104 hemoglobin seperti talasemia, HbS, dan HbC akan mengga
nggu pertumbuhan dan \line \up0 \expndtw0\charscalex102 replikasi merozoit. Seca
ra alami, defek eritrosit merupakan pertahanan terhadap \line \up0 \expndtw0\cha
rscalex103 penyakit malaria. Berikut ini akan diuraikan beberapa bentuk defek er
itrosit dan \line \up0 \expndtw-2\charscalex100 peranannya sebagai faktor resist
ensi terhadap parasit penyebab malaria. \par\pard\ql \li1800\sb0\sl-253\slmult0
\par\pard\ql\li1800\sb105\sl-253\slmult0 \up0 \expndtw-3\charscalex100 \ul0\nosu
persub\cf3\f4\fs22 1. Defisiensi Glucosa 6-Phosphat Dehidrogenase (G6PD) \par\pa

rd\qj \li1800\ri1625\sb4\sl-306\slmult0\fi720 \up0 \expndtw0\charscalex103 \ul0\

nosupersub\cf6\f7\fs22 Enzim G6PD terdiri dari polipeptida tetramer yang identik
, masing-masing \up0 \expndtw0\charscalex100 polipeptida tersusun dari 515 asam
amino. Gen G6PD terletak di bagian telomere \up0 \expndtw-1\charscalex100 l
engan panjang kromosom-x. Terdapat 4 macam variasi gen G6PD pada manusia, yaitu
\up0 \expndtw-2\charscalex100 Gd\ul0\super\cf7\f8\fs21 B\ul0\nosupersub\cf6\f7\f
s22 , Gd\ul0\super\cf7\f8\fs21 A \ul0\nosupersub\cf6\f7\fs22 , Gd\ul0\super\cf7\
f8\fs21 A-\ul0\nosupersub\cf6\f7\fs22 , dan Gd\ul0\super\cf7\f8\fs21 Med\ul0\nos
upersub\cf6\f7\fs22 . \par\pard\ql \li2159\sb58\sl-253\slmult0 \up0 \expndtw-2\c
harscalex100 a.\ul0\nosupersub\cf8\f9\fs22 \ul0\nosupersub\cf6\f7\fs22 Gen Gd
\ul0\super\cf7\f8\fs21 B\ul0\nosupersub\cf6\f7\fs22 (normal) \par\pard\li2159\s
b53\sl-253\slmult0\fi0\tx7660 \up0 \expndtw-1\charscalex100 b.\ul0\nosupersub\cf
8\f9\fs22 \ul0\nosupersub\cf6\f7\fs22 Gen Gd\ul0\super\cf7\f8\fs21 A\ul0\nosu
persub\cf6\f7\fs22 , terjadi substitusi nukleotida nomor\tab \up0 \expndtw-1
\charscalex100 367 (dari guanin menjadi\par\pard\li2159\sb54\sl-253\slmult0\f
i360\tx9424\tx9949 \up0 \expndtw-1\charscalex100 adenin), sehingga menyebabkan
perubahan asam amino nomor\tab \up0 \expndtw-1\charscalex100 126\tab \up0 \
expndtw-1\charscalex100 (dari\par\pard\li2159\sb53\sl-253\slmult0\fi360 \up0 \ex
pndtw-1\charscalex100 asparagin menjadi asam aspartat)\par\pard\qj \li2160\ri162
6\sb10\sl-300\slmult0\tx2520 \up0 \expndtw0\charscalex105 c.\ul0\nosupersub\cf8\
f9\fs22 \ul0\nosupersub\cf6\f7\fs22 Gen Gd\ul0\super\cf7\f8\fs21 A-\ul0\nosup
ersub\cf6\f7\fs22 , terjadi substitusi nukleotida nomor 2027 (dari adenin menjad
i \line\tab \up0 \expndtw0\charscalex101 guanin), sehingga mengubah asam ami
no nomor \up0 \expndtw-3\charscalex100 67 \up0 \expndtw0\charscalex101 (dari v
alin menjadi \par\pard\qj \li2520\ri1624\sb0\sl-310\slmult0 \up0 \expndtw0\char
scalex104 metionin). Pada gen ini juga terjadi substitusi nukleotida nomor 367 (
dari \up0 \expndtw0\charscalex100 guanin menjadi adenin), sehingga menyebabkan p
erubahan asam amino monor \up0 \expndtw-3\charscalex100 126 (dari asparagin menj
adi asam aspartat). \par\pard\li2160\sb49\sl-253\slmult0\fi0 \up0 \expndtw0\char
scalex103 d.\ul0\nosupersub\cf8\f9\fs22 \ul0\nosupersub\cf6\f7\fs22 Gen Gd\ul0
\super\cf7\f8\fs21 med\ul0\nosupersub\cf6\f7\fs22 , terjadi substitusi nukleotid
a nomor 563 (dari sitosin menjadi\par\pard\li2160\sb53\sl-253\slmult0\fi359\tx7
769\tx8304 \up0 \expndtw0\charscalex102 timin), sehingga mengubah asam amino
nomor\tab \up0 \expndtw0\charscalex102 188\tab \up0 \expndtw0\charscalex102 (d
ari serin menjadi\par\pard\li2160\sb54\sl-253\slmult0\fi359 \up0 \expndtw0\cha
rscalex102 fenilalanin)\par\pard\sect\sectd\fs24\paperw12240\paperh15840\pard\sb
0\sl-240{\bkmkstart Pg7}{\bkmkend Pg7}\par\pard\ql \li10320\sb0\sl-276\slmult0 \
par\pard\ql\li10320\sb195\sl-276\slmult0 \up0 \expndtw-3\charscalex100 \ul0\nosu
persub\cf1\f2\fs24 7 \par\pard\qj \li1799\sb0\sl-305\slmult0 \par\pard\qj\li1799
\ri1625\sb136\sl-305\slmult0\fi0 \up0 \expndtw0\charscalex105 \ul0\nosupersub\cf
6\f7\fs22 Enzim G6PD yang dibawa gen Gd\ul0\super\cf7\f8\fs21 B\ul0\nosupersub\c
f6\f7\fs22 dan Gd\ul0\super\cf7\f8\fs21 A\ul0\nosupersub\cf6\f7\fs22 mempunyai
aktivitas normal, gen Gd\ul0\super\cf7\f8\fs21 A-\line \up0 \expndtw0\charscale
x103 \ul0\nosupersub\cf6\f7\fs22 mempunyai aktivitas normal 8-20% dan Gd\ul0\sup
er\cf7\f8\fs21 med\ul0\nosupersub\cf6\f7\fs22 hanya 5% yang beraktivitas normal
. \line \up0 \expndtw0\charscalex100 Secara normal, G6PD mempunyai half life 62
hari, sehingga dapat mempertahankan \line \up0 \expndtw0\charscalex104 GSH selam
a 100-120 hari (life span eritrosit). Pada individu yang mempunyai gen \line \up
0 \expndtw-1\charscalex100 Gd\ul0\super\cf7\f8\fs21 A-\ul0\nosupersub\cf6\f7\fs2
2 , enzim G6PD mempunyai half life hanya \up0 \expndtw-1\charscalex100 13
hari, sehingga eritrosit akan \par\pard\qj \li1799\ri1627\sb0\sl-320\slmult0
\up0 \expndtw0\charscalex100 mengalami kerusakan secara prematur ({\field{\*\fl
dinst {HYPERLINK "http://www.as.ua.edu/ant/bindon/ant475/" }}{\fldrslt {\ul0\nos
upersub\cf12\f13\fs22\ul http://www.as.ua.edu/ant/bindon/ant475/}}} \line \up0 \
expndtw-3\charscalex100 \ul0\nosupersub\cf12\f13\fs22\ul g6pd/img003.jpg\ul0\nos
upersub\cf6\f7\fs22 ). \par\pard\qj \li1799\ri1625\sb0\sl-300\slmult0\fi720 \up0
\expndtw0\charscalex102 Defisiensi enzim G6PD merupakan faktor resistensi alami
terhadap malaria \line \up0 \expndtw0\charscalex103 (Ruwende et al., \up0 \exp
ndtw0\charscalex108 1995; Tishkoff et al., \up0 \expndtw0\charscalex103 2001). D
efisiensi G6PD mengakibatkan \par\pard\qj \li1800\ri1623\sb0\sl-305\slmult0\fi0

\up0 \expndtw0\charscalex104 pembentukan NADPH mengalami hambatan, sehingga deto

ksifikasi ROS menjadi \line \up0 \expndtw0\charscalex100 molekul non toksik a
kibat defisiensi NADPH menyebabkan eritrosit mengalami \line \up0 \expndtw0
\charscalex103 stress oksidatif sehingga mudah lisis (Marks et al., 2001). Tingg
inya produk ROS \line \up0 \expndtw0\charscalex104 pada penderita defisiensi G6P
D, selain menghancurkan eritrosit secara dini juga \line \up0 \expndtw-1\charsca
lex100 akan menghancurkan merozoit yang berada di dalamnya \up0 \expndtw-1
\charscalex100 (Janney, et al., \up0 \expndtw-3\charscalex100 1986; \par\pard\
qj \li1799\ri1626\sb0\sl-320\slmult0\fi0 \up0 \expndtw0\charscalex105 Friedman,
1979). Disebutkan juga bahwa P. falciparum tumbuh sangat lambat di \up0 \expndtw
-3\charscalex100 dalam eritrosit yang defisien G6PD (Roth et al., 1983). \par\pa
rd\ql \li1800\sb0\sl-253\slmult0 \par\pard\ql\li1800\sb62\sl-253\slmult0 \up0 \e
xpndtw-3\charscalex100 \ul0\nosupersub\cf3\f4\fs22 2. Talasemia \par\pard\qj \li
1800\ri1624\sb29\sl-300\slmult0\fi720 \up0 \expndtw0\charscalex100 \ul0\nosupers
ub\cf6\f7\fs22 Talasemia yaitu penyakit herediter yang disebabkan oleh kekuranga
n rantai \line \up0 \expndtw-1\charscalex100 globin pembentuk hemoglobin, baik r
antai globin \u945? (talasemia \u945?) maupun rantai globin \line \up0 \expndtw3\charscalex100 \u946? \up0 \expndtw0\charscalex101 (talasemia \u946?) \up0 \ex
pndtw-3\charscalex100 (Suryohudoyo, \up0 \expndtw0\charscalex102 2000). Talasem
ia disebutkan juga merupakan \par\pard\qj \li1799\ri1625\sb16\sl-305\slmult0\
fi0 \up0 \expndtw0\charscalex100 pertahanan alami terhadap malaria. Ketidakseimb
angan rantai globin pada talasemia \line \up0 \expndtw0\charscalex103 menyebabka
n oksidasi membran oleh hemikrom dan molekul-molekul derivat ROS \line \up0 \exp
ndtw0\charscalex100 (Grinberg et al., 1995; Sorensen et al., 1990). ROS, selain
menghancurkan membran \line \up0 \expndtw0\charscalex102 eritrosit juga menghanc
urkan merozoit di dalamnya (Clark et al., 1989). Secara in \line \up0 \expndtw0\
charscalex102 vitro, eritrosit dengan hemoglobin H sangat toksik terhadap merozo
it (Ifediba et \line \up0 \expndtw0\charscalex100 al., 1985; Yathavong et al., 1
988). Hemoglobin H dapat terjadi pada manusia dengan \line \up0 \expndtw-1\chars
calex100 delesi 3 gen \u945? talasemia (Zhu et al., 1993) dan delesi 2 gen \u945
? talasemia (Derry et al., \line \up0 \expndtw0\charscalex102 1988). Alpha tal
asemia merupakan pertahanan terhadap malaria dengan jalan \line \up0 \expn
dtw0\charscalex100 mengganggu respon imun terhadap eritrosit yang mengandung par
asit malaria (Luzzi \line \up0 \expndtw0\charscalex100 et al., 1991). Eritrosit
talasemia yang terinfeksi P. falciparum, kemampuannya untuk \line \up0 \expndtw0
\charscalex102 menempel pada sel endotel lebih rendah dibanding eritrosit normal
(Carlson et al., \line \up0 \expndtw-3\charscalex100 1994). \par\pard\ql \li179
9\sb0\sl-253\slmult0 \par\pard\ql\li1799\sb105\sl-253\slmult0 \up0 \expndtw-3\ch
arscalex100 \ul0\nosupersub\cf3\f4\fs22 3. Eliptositosis/Ovalosititosis \par\par
d\qj \li1799\ri1625\sb5\sl-305\slmult0\fi720 \up0 \expndtw0\charscalex100 \ul0\n
osupersub\cf6\f7\fs22 Ovalosit adalah kelainan herediter yang ditandai dengan er
itrosit berbentuk \line \up0 \expndtw-3\charscalex100 oval \up0 \expndtw0\charsc
alex106 (elip), terutama terdapat di Asia Tenggara. Kelainan ini diturunkan seca
ra \line \up0 \expndtw0\charscalex100 autosom dominan (Castelio et al. (1981) in
Suryohudoyo,2000). Ciri khas ovalositosis \line \up0 \expndtw0\charscalex104 ad
alah kekakuan eritrosit yang berbentuk oval, namun tanpa disertai anemia dan \li
ne \up0 \expndtw0\charscalex100 hemolisis. Percobaan in vitro menunjukkan bahwa
ovalositosis tahan terhadap invasi \par\pard\qj \li1800\ri1625\sb20\sl-300\slmul
t0 \up0 \expndtw0\charscalex100 P. falciparum (Kidson et al. (1981) in Suryohudo
yo, 2000) dan P. knowlesi (Hardly et \up0 \expndtw-3\charscalex100 al. (1983) in
Suryohudoyo, 2000). \par\pard\sect\sectd\fs24\paperw12240\paperh15840\pard\sb0\
sl-240{\bkmkstart Pg8}{\bkmkend Pg8}\par\pard\ql \li10320\sb0\sl-276\slmult0 \pa
r\pard\ql\li10320\sb195\sl-276\slmult0 \up0 \expndtw-3\charscalex100 \ul0\nosupe
rsub\cf1\f2\fs24 8 \par\pard\qj \li1799\sb0\sl-304\slmult0 \par\pard\qj\li1799\r
i1625\sb137\sl-304\slmult0\fi720 \up0 \expndtw0\charscalex103 \ul0\nosupersub\cf
6\f7\fs22 Secara seluler diketahui bahwa eritrosit ovalosit mempunyai kelainan p
ada \up0 \expndtw0\charscalex100 protein band-3 (sebagai anion chanel), yaitu pr
otein transmembran, dengan ujung N \up0 \expndtw-2\charscalex100 terletak di sis
i sitoplasma dan berfungsi sebagai saluran yang mengalirkan ion Cl\ul0\super\cf7
\f8\fs21 -\ul0\nosupersub\cf6\f7\fs22 dan \up0 \expndtw0\charscalex100 HCO\ul0\

sub\cf7\f8\fs21 3-\ul0\nosupersub\cf6\f7\fs22 . Daerah sitoplasma protein ba

nd-3 terkait dengan sitoskeleton melalui \up0 \expndtw0\charscalex100 protei
n ankirin dan spektrin B (Suryohudoyo, 2000). Penelitian menunjukkan bahwa \up0
\expndtw-3\charscalex100 ovalositosis mengalami perubahan protein band-3 sebagai
berikut: \par\pard\ql \li2519\ri1625\sb20\sl-300\slmult0\tx2880\tx2880 \up0 \ex
pndtw0\charscalex105 1.\ul0\nosupersub\cf8\f9\fs22 \ul0\nosupersub\cf6\f7\fs22
mutasi titik pada nukleotida nomor 166 (dari adenin menjadi guanine), \line\ta
b \up0 \expndtw-1\charscalex100 sehingga menghasilkan perubahna asam amino nomor
58 (dari lisin menjadi \line\tab \up0 \expndtw-3\charscalex100 glutamate) \up0
\expndtw0\charscalex100 (Jarolim et al. \up0 \expndtw-1\charscalex100 (1991)
in Suryohudoyo, \up0 \expndtw0\charscalex100 2000). Mutasi ini \par\pard\qj \
li2880\ri1626\sb20\sl-300\slmult0\fi0 \up0 \expndtw-1\charscalex100 disebutkan h
anya merupakan bentuk polimorfisme protein band-3, karena \up0 \expndtw-3\charsc
alex100 juga ditemukan pada eritrosit normal. \par\pard\ql \li2520\ri1624\sb0\sl
-320\slmult0\tx2880 \up0 \expndtw0\charscalex100 2.\ul0\nosupersub\cf8\f9\fs22
\ul0\nosupersub\cf6\f7\fs22 delesi nukleotida nomor 1190-1224 yang mengakibatka
n hilangnya 9 asam \line\tab \up0 \expndtw-3\charscalex100 amino (Jarolim et al.
(1991) in Suryohudoyo, 2000). \par\pard\ql \li2520\ri1623\sb0\sl-305\slmult0\tx
2880\tx2880\tx2880\tx2880 \up0 \expndtw0\charscalex102 3.\ul0\nosupersub\cf8\f9\
fs22 \ul0\nosupersub\cf6\f7\fs22 ada tambahan 13 asam amino pada protein band3 (Jones et al. (1991) in \line\tab \up0 \expndtw0\charscalex101 Suryohudoyo, 20
00). Belum diketahui dengan pasti apakah tambahan 13 \line\tab \up0 \expndtw0\ch
arscalex101 asam amino tersebut murni merupakan mutasi protein band-3 at
au \line\tab \up0 \expndtw0\charscalex100 kontaminan dari parasit malaria, karen
a ke-13 asam amino tersebut mirip \line\tab \up0 \expndtw0\charscalex102 dengan
protein ring infected erythrocyte surface antigen \up0 \expndtw-2\charscal
ex100 (RESA) \par\pard\ql \li2880\sb35\sl-253\slmult0 \up0 \expndtw-3\charscalex
100 (Christy dan Ruff, (1991) in Suryohudoyo, 2000). \par\pard\qj \li1800\ri1625
\sb304\sl-306\slmult0\fi719 \up0 \expndtw0\charscalex105 Ovalositosis dapat memp
roteksi penderitanya dari invasi parasit malaria. \line \up0 \expndtw0\charscale
x103 Proteksi tersebut berhubungan dengan kekakuan membran ovalosit, sehi
ngga \line \up0 \expndtw-1\charscalex100 menghambat proses invaginasi dan
endositosis merozoit ke dalam eritrosit \line \up0 \expndtw0\charscale
x102 (Mohandas et al. \up0 \expndtw0\charscalex101 (1984) in Suryohudoyo, \u
p0 \expndtw0\charscalex102 2000). Delesi asam amino protein \par\pard\qj \li
1800\ri1605\sb14\sl-306\slmult0 \up0 \expndtw0\charscalex102 transmembran band-3
menyebabkan ikatan yang lebih erat dengan protein ankirin \line \up0 \expndtw0\
charscalex105 sehingga mengurangi mobilitas lateral membran. Eratnya ikatan prot
ein band-3-\line \up0 \expndtw0\charscalex105 ankirin menyebabkan hambatan prose
s invaginasi, karena protein tersebut tidak \line \up0 \expndtw0\charscalex102 d
apat didorong keluar dari daerah tempat pengikatan merozoit. Dijelaskan bahwa \l
ine \up0 \expndtw0\charscalex100 ikatan merozoit-eritrosit akan diikuti peru
bahan drastis komponen proteinnya, \line \up0 \expndtw-2\charscalex100 sehing
ga pada daerah kontak antara membran eritrosit-merozoit tidak la
gi \line \up0 \expndtw-2\charscalex100 mengandung protein band-3 (Dluzewski et a
l. (1989) in Suryohudoyo, 2000). \par\pard\qj \li1799\ri1625\sb0\sl-305\slmult0\
fi720 \up0 \expndtw0\charscalex101 Kekakuan membran bukan satu-satunya penyebab
hambatan invasi merozoit \up0 \expndtw0\charscalex100 pada ovalosit (Dluzewski e
t al. (1992) in Suryohudoyo, 2000). Rendahnya kadar ATP \up0 \expndtw-1\charscal
ex100 sel ovalosit dibandingkan eritrosit normal, menyebabkan merozoit tidak dap
at masuk \up0 \expndtw0\charscalex108 sel. Merozoit tidak dapat memasuki sel jik
a kadar ATP intrasel di bawah nilai \up0 \expndtw0\charscalex103 ambang tertentu
yaitu 0,11mM. Rendahnya kadar ATP pada ovalosit, kemungkinan \up0 \expndtw0\cha
rscalex100 disebabkan oleh gangguan saluran anion, sehingga pengeluaran HCO\ul0\
sub\cf7\f8\fs21 3-\ul0\nosupersub\cf6\f7\fs22 terganggu dan \up0 \expndtw0\char
scalex100 menyebabkan pH ovalosit meningkat. Peningkatan pH selanjutnya akan men
gganggu \up0 \expndtw0\charscalex100 enzim-enzim tertentu yang terlibat dala
m proses glikolisis, yaitu satu-satunya \up0 \expndtw-3\charscalex100 sumber
ATP bagi eritrosit (Suryohudoyo, 2000). \par\pard\ql \li1800\sb0\sl-253\slmult0
\par\pard\ql\li1800\sb100\sl-253\slmult0 \up0 \expndtw-1\charscalex100 \ul0\nos

upersub\cf3\f4\fs22 4.\ul0\nosupersub\cf9\f10\fs22 \ul0\nosupersub\cf3\f4\fs22

Tidak Mempunyai Antigen Duffy \par\pard\sect\sectd\fs24\paperw12240\paperh15840\
pard\sb0\sl-240{\bkmkstart Pg9}{\bkmkend Pg9}\par\pard\ql \li10320\sb0\sl-276\sl
mult0 \par\pard\ql\li10320\sb195\sl-276\slmult0 \up0 \expndtw-3\charscalex100 \u
l0\nosupersub\cf1\f2\fs24 9 \par\pard\qj \li1799\sb0\sl-300\slmult0 \par\pard\qj
\li1799\ri1627\sb145\sl-300\slmult0\fi720 \up0 \expndtw0\charscalex100 \ul0\nosu
persub\cf6\f7\fs22 Antigen Duffy pada membran eritrosit digunakan sebagai perant
ara P. vivax \line \up0 \expndtw0\charscalex103 dan P. knowlesi masuk ke da
lam eritrosit \up0 \expndtw0\charscalex102 (Gelpi and King, \up0 \expndtw0\ch
arscalex100 1976). Merozoit \par\pard\qj \li1799\ri1624\sb0\sl-310\slmult0 \up0
\expndtw0\charscalex105 Plasmodium tersebut mempunyai protein Duffy binding pro
tein (DBP), ligan dari \line \up0 \expndtw-1\charscalex100 reseptor Duffy pa
da eritrosit. Ikatan antigen Duffy-DBP berperan dalam \line \up0 \ex
pndtw0\charscalex101 mekanisme invasi, terutama dalam pembentukan tight ju
nction \up0 \expndtw-3\charscalex100 (Reed, \up0 \expndtw-3\charscalex100 2000).
\par\pard\qj \li1800\ri1624\sb0\sl-310\slmult0\fi0 \up0 \expndtw0\charscalex100
Ketiadaan antigen Duffy akan menurunkan kekuatan intraksi merozoit-eritr
osit \up0 \expndtw-1\charscalex100 sehingga invasi gagal dilakukan. Manusia yang
mempunyai abnormalitas antigen duffy, \up0 \expndtw0\charscalex104 menunjukkan
resisten terhadap P. vivax (Hamblin dan DiRienzo, 2000). Di Afrika \up0 \expndtw
0\charscalex100 barat tidak banyak terjadi infeksi oleh P. vivax, karena sebagia
n besar penduduk di \up0 \expndtw-2\charscalex100 daerah ini eritrositnya tidak
mengandung antigen duffy (Wakelin, 1988). \par\pard\ql \li1800\sb0\sl-253\slmult
0 \par\pard\ql\li1800\sb67\sl-253\slmult0 \up0 \expndtw-1\charscalex100 \ul0\nos
upersub\cf3\f4\fs22 5.\ul0\nosupersub\cf9\f10\fs22 \ul0\nosupersub\cf3\f4\fs22
Sickle Cell (Sel Bulan Sabit)/ HbS \par\pard\qj \li1799\ri1624\sb25\sl-305\slmul
t0\fi720 \up0 \expndtw0\charscalex105 \ul0\nosupersub\cf6\f7\fs22 Eritrosit yang
mengandung hemoglobin S (Sickle) berbentuk bulan sabit, \line \up0 \expndtw0\ch
arscalex100 sehingga disebut sel bulan sabit. Secara epidemiologis terlihat bahw
a distribusi gen \line \up0 \expndtw0\charscalex100 HbS banyak terdapat di d
aerah endemik malaria. Manusia yang terinfeksi P. \line \up0 \expndtw-1\ch
arscalex100 falciparum nampak lebih survive terhadap infeksi akut pada mereka ya
ng mempunyai \line \up0 \expndtw0\charscalex104 sel bulan sabit dari pada e
ritrosit normal. Manusia dengan homozigot HbS \line \up0 \expndtw0\charscal
ex106 (HbS/HbS) rentan terhadap kematian akibat komplikasi antara malaria dengan
\line \up0 \expndtw0\charscalex103 penyakit sickle cell. Pada penderita heteroz
igot HbS (HbA/HbS) tahan terhadap \line \up0 \expndtw0\charscalex106 malaria, ka
rena tidak ada komplikasi dengan keberadaan gen HbS. Anak dengan \line \up0 \exp
ndtw-1\charscalex100 HbS/HbA survive terhadap infeksi akut malaria dibanding HbA
/HbA (Ringelhann et \line \up0 \expndtw-3\charscalex100 al., \up0 \expndtw0\char
scalex102 1976). Dapat disimpulkan bahwa penderita sel bulan sabit yang tahan te
rhadap \line \up0 \expndtw0\charscalex103 malaria adalah heterozigot HbS (HBA/Hb
S), dan bukan penderita sel bulan sabit \line \up0 \expndtw-3\charscalex100 homo
zigot (HbS/HbS) \par\pard\qj \li1799\ri1625\sb15\sl-306\slmult0\fi720 \up0 \expn
dtw0\charscalex100 Eritrosit pada heterozigot HbS tidak menunjukkan bentuk sel s
eperti bulan \line \up0 \expndtw0\charscalex102 sabit jika tekanan oksigen dalam
darahnya normal, dan baru berbentuk bulan sabit \line \up0 \expndtw0\charscalex
100 jika terjadi penurunan tekanan oksigen dalam darah. Pada manusia heterozigot
HbS \line \up0 \expndtw-1\charscalex100 yang melakukan aktivitas berlebihan, me
nyebabkan jumlah sel bulan sabit mengalami \line \up0 \expndtw-1\charscalex100 p
eningkatan (Martin et al., 1989). Sel heterozigot HbS yang terinfeksi P. falcipa
rum \line \up0 \expndtw0\charscalex103 akan mengalami deformasi menjadi sel
bulan sabit, karena parasit tersebut \line \up0 \expndtw0\charscalex102 men
yebabkan penurunan tekanan oksigen dalam eritrosit sampai di bawah ambang. \line
\up0 \expndtw0\charscalex100 Deformasi eritrosit tersebut merupakan tanda
bahwa sel tersebut abnormal, \line \up0 \expndtw0\charscalex104 sehingga menj
adi target sel-sel fagosit untuk didestruksi (Luzzato et al., 1970). \line \up0
\expndtw0\charscalex103 Penelitian Anastasi (1984) menunjukkan bahwa pembentukan
ROS dalam sel bulan \line \up0 \expndtw0\charscalex100 sabit menyebabkan hamb
atan pertumbuhan P. falciparum dan kematian parasit \line \up0 \expndtw0\c

harscalex100 tersebut. Sel bulan sabit menghasilkan lebih banyak ROS seperti sup
eroxide anion \line \up0 \expndtw0\charscalex106 (O\ul0\sub\cf7\f8\fs21 2-\ul0\n
osupersub\cf6\f7\fs22 ) dan hidrogen peroksida (H\ul0\sub\cf7\f8\fs21 2\ul0\nosu
persub\cf6\f7\fs22 O\ul0\sub\cf7\f8\fs21 2\ul0\nosupersub\cf6\f7\fs22 ) dibandin
gkan eritrosit normal (Rank et al., \line \up0 \expndtw-3\charscalex100 1985). \
par\pard\ql \li1799\sb38\sl-253\slmult0 \up0 \expndtw0\charscalex100 \ul0\nosupe
rsub\cf3\f4\fs22 6.\ul0\nosupersub\cf9\f10\fs22 \ul0\nosupersub\cf3\f4\fs22 Hem
oglobin C (HbC) \par\pard\qj \li1799\ri1626\sb0\sl-310\slmult0\fi720 \up0 \expnd
tw0\charscalex100 \ul0\nosupersub\cf6\f7\fs22 Hemoglobin C disebutkan dapa
t menghambat infeksi malaria. Studi \line \up0 \expndtw0\charscalex100 e
pidemiologis menunjukkan bahwa manusia dengan homozigot atau heterozigot HbC \li
ne \up0 \expndtw0\charscalex100 tahan terhadap infeksi parasit malaria (Willcox
et al., 1983). P. falciparum secara \par\pard\sect\sectd\fs24\paperw12240\paperh
15840\pard\sb0\sl-240{\bkmkstart Pg10}{\bkmkend Pg10}\par\pard\ql \li10200\sb0\s
l-276\slmult0 \par\pard\ql\li10200\sb195\sl-276\slmult0 \up0 \expndtw-3\charscal
ex100 \ul0\nosupersub\cf1\f2\fs24 10 \par\pard\qj \li1800\sb0\sl-304\slmult0 \pa
r\pard\qj\li1800\ri1624\sb137\sl-304\slmult0 \up0 \expndtw0\charscalex100 \ul0\n
osupersub\cf6\f7\fs22 signifikan lebih rendah menyerang manusia heterozigot HbC
dibanding HbA. Namun \line \up0 \expndtw0\charscalex104 jika dibandingkan, resik
o malaria lebih rendah pada homozigot HbC dibandingkan \line \up0 \expndtw-1\cha
rscalex100 heterozigot HbC. Homozigot HbC yang terserang malaria hanya menunjukk
an anemia \line \up0 \expndtw0\charscalex105 dan splenomegali ringan (Modiano et
al., 2001). Mekanisme penghambatan invasi \line \up0 \expndtw0\charscalex103 Pl
asmodium sp. pada HbC mungkin berhubungan dengan defek pengikatan oksigen \line
\up0 \expndtw-3\charscalex100 oleh eritrosit. \par\pard\ql \li1800\sb0\sl-253\sl
mult0 \par\pard\ql\li1800\sb106\sl-253\slmult0 \up0 \expndtw-3\charscalex100 \ul
0\nosupersub\cf3\f4\fs22 Kesimpulan \par\pard\qj \li1800\ri1624\sb4\sl-306\slmul
t0\fi720 \up0 \expndtw0\charscalex100 \ul0\nosupersub\cf6\f7\fs22 Keberhasilan m
ekanisme invasi dan pertumbuhan merozoit sangat ditentukan \up0 \expndtw0\charsc
alex104 oleh kondisi eritrosit sebagai habitatnya. Mekanisme invasi melibatkan i
nteraksi \up0 \expndtw0\charscalex100 membran eritrosit dan membran merozoit, se
hingga defek pada membran eritrosit \up0 \expndtw0\charscalex104 seperti defisi
ensi glikoporin, antigen Duffy dan defek protein band-3 akan \up0 \expnd
tw0\charscalex100 menyebabkan berkurangnya kekuatan interaksi dan inva
si gagal dilakukan. \up0 \expndtw0\charscalex104 Pertumbuhan dan perkemban
gan merozoit selama di dalam eritrosit, sangat \up0 \expndtw0\charscalex10
6 ditentukan oleh kondisi sitoplasma eritrosit. Jika di dalam sitoplasma eritros
it \up0 \expndtw0\charscalex100 mengandung faktor-faktor yang memberi suasan
a tidak nyaman bagi merozoit, \up0 \expndtw0\charscalex100 seperti defisiens
i enzim G6PD, dan mutasi hemoglobin seperti talasemia, HbS, dan \up0 \expndtw-3\
charscalex100 HbC akan mengganggu pertumbuhan dan replikasi merozoit. \par\pard\
ql \li1800\sb0\sl-253\slmult0 \par\pard\ql\li1800\sb0\sl-253\slmult0 \par\pard\q
l\li1800\sb152\sl-253\slmult0 \up0 \expndtw-3\charscalex100 \ul0\nosupersub\cf3\
f4\fs22 Daftar Pustaka \par\pard\ql \li1800\sb0\sl-310\slmult0 \par\pard\ql\li18
00\ri1625\sb10\sl-310\slmult0\tx2340\tx2340 \up0 \expndtw0\charscalex104 \ul0\no
supersub\cf6\f7\fs22 Anastasi, J. 1984. Hemoglobin S-mediated membrane oxidant i
njury: protection \line\tab \up0 \expndtw0\charscalex103 from malaria and pathol
ogy in sickle cell disease. Med. Hypotheses. 14: 311-\line \tab \up0 \expndtw-3\
charscalex100 320 \par\pard\ql \li1800\ri1625\sb290\sl-310\slmult0\tx2340\tx2339
\up0 \expndtw0\charscalex104 Carlson, J., Nash, G.B., Gabutti, V., al-Yaman F.,
and Wahlgren, M. 1994. Natural \line\tab \up0 \expndtw0\charscalex100 protectio
n against severe Plasmodium falciparum malaria due to impaired \line\tab
\up0 \expndtw-3\charscalex100 rosette formation. Blood. 84:3909-3814. \par\pard
\qj \li1800\sb0\sl-300\slmult0 \par\pard\qj\li1800\ri1625\sb19\sl-300\slmult0\tx
2339 \up0 \expndtw0\charscalex105 Clark, I.A., Chaudhri, G., and Cowden, W.B. 19
89. Some roles of free radicals in \line\tab \up0 \expndtw-3\charscalex100 malar
ia. Free. Radic. Biol. Med. 6: 315-321. \par\pard\qj \li1799\ri1625\sb284\sl-320
\slmult0\tx2339 \up0 \expndtw0\charscalex100 Derry, S., Wood, W.G., Pippard, M.,
Clegg, J.B., Weatherall, D.J., Wickramasinghe, \line\tab \up0 \expndtw0\charsca
lex106 S.N., Darley, J., Fucharoen, S., and Wasi, P. \up0 \expndtw-3\char

scalex100 1988. Hematologic and \par\pard\qj \li2339\ri1627\sb0\sl-300\slmul

t0 \up0 \expndtw0\charscalex100 biosynthetic studies in homozygous hemoglobi
n Constant Spring. J. Clin. \up0 \expndtw-3\charscalex100 Invest. 173:1673-1
682. \par\pard\qj \li1800\sb0\sl-300\slmult0 \par\pard\qj\li1800\ri1993\sb17\sl300\slmult0\tx2340 \up0 \expndtw-2\charscalex100 Fahmi, U. 2001. Malaria di Temp
at Pengungsian. Partnership Meeting In Roll Back \line\tab \up0 \expndtw-3\chars
calex100 Malaria. Jakarta. \par\pard\sect\sectd\fs24\paperw12240\paperh15840\par
d\sb0\sl-240{\bkmkstart Pg11}{\bkmkend Pg11}\par\pard\ql \li10200\sb0\sl-276\slm
ult0 \par\pard\ql\li10200\sb195\sl-276\slmult0 \up0 \expndtw-3\charscalex100 \ul
0\nosupersub\cf1\f2\fs24 11 \par\pard\qj \li1800\sb0\sl-300\slmult0 \par\pard\qj
\li1800\ri1626\sb145\sl-300\slmult0\tx2339 \up0 \expndtw0\charscalex107 \ul0\nos
upersub\cf6\f7\fs22 Friedman, M.J. 1979. Oxidant damage mediates variant red cel
l resistance to \line\tab \up0 \expndtw-3\charscalex100 malaria. Nature. 280:245
-247 \par\pard\qj \li1800\sb0\sl-300\slmult0 \par\pard\qj\li1800\ri1626\sb20\sl300\slmult0\tx2340 \up0 \expndtw0\charscalex103 Gelpi, A.P.,and King, M.C. 1976.
Association of Duffy blood groups with the sickle \line\tab \up0 \expndtw-3\cha
rscalex100 cell trait. Hum. Genet. 32: 65-68. \par\pard\qj \li1799\sb0\sl-300\sl
mult0 \par\pard\qj\li1799\ri1628\sb20\sl-300\slmult0\tx2340 \up0 \expndtw-1\char
scalex100 Glucosa-6-phosphate dehydrogenase and Malaria. {\field{\*\fldinst {HYP
ERLINK "http://www.as.ua.edu/ant/bindon/" }}{\fldrslt {\ul0\nosupersub\cf12\f13\
fs22\ul http://www.as.ua.edu/ant/bindon/}}} \line \ul0\tab \up0 \expndtw-2\chars
calex100 \ul0\nosupersub\cf12\f13\fs22\ul ant475/g6pd/img003.jpg\ul0\nosupersub\
cf6\f7\fs22 . Dikunjungi 17 Oktober 2004 \par\pard\qj \li1800\sb0\sl-300\slmult0
\par\pard\qj\li1800\ri1625\sb20\sl-300\slmult0\tx2340 \up0 \expndtw0\charscalex
104 Gratzer, W.B., and Dluzewski, A.R. 1993. The red blood cell and malaria para
site \line\tab \up0 \expndtw-3\charscalex100 invasion. Semin. Hematol. 30: 232-2
47 \par\pard\ql \li1800\sb0\sl-300\slmult0 \par\pard\ql\li1800\ri1625\sb20\sl-30
0\slmult0\tx2340\tx2340 \up0 \expndtw0\charscalex103 Grinberg, L.N., Rachmilewit
z, E.A., Kitrossky, N., and Chevion, M. 1995. Hydroxyl \line\tab \up0 \expndtw0\
charscalex102 radical generation in beta-thalassemic red blood cells. Free Radic
. Biol. Med. \line\tab \up0 \expndtw-3\charscalex100 18:611-615 \par\pard\ql \li
1800\sb0\sl-253\slmult0 \par\pard\ql\li1800\sb106\sl-253\slmult0\tx5533 \up0 \ex
pndtw0\charscalex108 Hamblin, M.T., and Di Rienzo, A. \tab \up0 \expndtw0\charsc
alex107 2000. Detection of the signature of natural \par\pard\qj \li2340\ri1625\
sb0\sl-320\slmult0 \up0 \expndtw0\charscalex102 selection in humans: evidence fr
om the Duffy blood group locus. Am. J. Hum. \up0 \expndtw-3\charscalex100 Genet.
66: 1669-1679. \par\pard\qj \li1799\ri2106\sb272\sl-320\slmult0\tx2340 \up0 \ex
pndtw-2\charscalex100 Hemoglobin.{\field{\*\fldinst {HYPERLINK "http://www.brain
yencyclopecia.com/encyclopedia/h/he/hemoglobin./" }}{\fldrslt {\ul0\nosupersub\c
f12\f13\fs22\ul http://www.brainyencyclopecia.com/encyclopedia/h/he/hemoglobin.}
}} \line \ul0\tab \up0 \expndtw-3\charscalex100 \ul0\nosupersub\cf12\f13\fs22\ul
html\ul0\nosupersub\cf6\f7\fs22 . Dikunjungi 17 Oktober 2004. \par\pard\ql \li1
799\sb0\sl-253\slmult0 \par\pard\ql\li1799\sb83\sl-253\slmult0 \up0 \expndtw-2\c
harscalex100 Hyde, J. E. 1990. Molecular Parasitology. Van Nostrand Reinhold. Ne
w York. \par\pard\ql \li1799\sb0\sl-300\slmult0 \par\pard\ql\li1799\ri1627\sb29\
sl-300\slmult0\tx2339\tx2339 \up0 \expndtw-2\charscalex100 Ifediba, T.C., Stern,
A., Ibrahim, A., and Rieder, R.F. 1985. Plasmodium falciparum in \line\tab \up0
\expndtw-1\charscalex100 vitro: diminished growth in hemoglobin H disease eryth
rocytes. Blood. 65:452-\line \tab \up0 \expndtw-2\charscalex100 455. \par\pard\q
l \li1800\sb0\sl-310\slmult0 \par\pard\ql\li1800\ri1624\sb2\sl-310\slmult0\tx234
0\tx2340 \up0 \expndtw-2\charscalex100 Janney, S.K., Joist, J.J., and Fitch, C.D
. 1986. Excess release of ferriheme in G6PD-\line \tab \up0 \expndtw0\charscalex
100 deficient erythrocytes: possible cause of hemolysis and resistance to malari
a. \line\tab \up0 \expndtw-3\charscalex100 Blood. 67:331-333 \par\pard\ql \li180
0\sb0\sl-253\slmult0 \par\pard\ql\li1800\sb85\sl-253\slmult0 \up0 \expndtw0\char
scalex100 Koolman, J. dan Rohm, K.H. 2001. Atlas berwarna dan teks biokimia. Ali
h bahasa: S. \par\pard\ql \li2339\sb67\sl-253\slmult0 \up0 \expndtw-3\charscalex
100 I. Wanandi. Hipokrates. Jakarta. \par\pard\li1800\sb0\sl-264\slmult0\par\par
d\li1800\sb102\sl-264\slmult0\fi0\tx5312 \up0 \expndtw0\charscalex104 Life cycl
e of Plasmodium sp.\tab \up0 \expndtw-2\charscalex100 2004. \ul0\nosupersub\c

f1\f2\fs24 {\field{\*\fldinst {HYPERLINK "http://whyfiles.org/016skeeter/malari

a2.html./" }}{\fldrslt {\ul0\nosupersub\cf11\f12\fs24\ul http://whyfiles.org/016
skeeter/malaria2.html\ul0\nosupersub\cf1\f2\fs24 .}}}\par\pard\li1800\sb3\sl-276
\slmult0\fi539 \up0 \expndtw-3\charscalex100 \ul0\nosupersub\cf1\f2\fs24 Dikunju
ngi 7 Oktober 2004\par\pard\ql \li1800\ri1624\sb292\sl-310\slmult0\tx2339\tx2339
\up0 \expndtw0\charscalex104 \ul0\nosupersub\cf6\f7\fs22 Luzzatto, L., Nwachuku
-Jarrett, E.S., and Reddy, S. 1970. Increased sickling of \line\tab \up0 \expndt
w0\charscalex104 parasitised erythrocytes as mechanism of resistance against mal
aria in the \line\tab \up0 \expndtw-3\charscalex100 sickle-cell trait. Lancet 1.
7642:319-321. \par\pard\sect\sectd\fs24\paperw12240\paperh15840\pard\sb0\sl-240
{\bkmkstart Pg12}{\bkmkend Pg12}\par\pard\ql \li10200\sb0\sl-276\slmult0 \par\pa
rd\ql\li10200\sb195\sl-276\slmult0 \up0 \expndtw-3\charscalex100 \ul0\nosupersub
\cf1\f2\fs24 12 \par\pard\ql \li1800\sb0\sl-306\slmult0 \par\pard\ql\li1800\sb0\
sl-306\slmult0 \par\pard\ql\li1800\ri1624\sb128\sl-306\slmult0\tx2339\tx2340\tx2
340 \up0 \expndtw0\charscalex102 \ul0\nosupersub\cf6\f7\fs22 Luzzi, G.A., Merry,
A.H., Newbold, C.I., Marsh, K., Pasvol, G., and Weatherall, D.J. \line\tab \up0
\expndtw0\charscalex102 1991. Surface antigen expression on Plasmodiu
m falciparum-infected \line\tab \up0 \expndtw0\charscalex105 erythrocytes is m
odified in alpha- and beta-thalassemia. J. Exp. Med. 173: \line\tab \up0 \expndt
w-3\charscalex100 785-791 \par\pard\qj \li1800\sb0\sl-300\slmult0 \par\pard\qj\l
i1800\ri1624\sb19\sl-300\slmult0\tx2520 \up0 \expndtw0\charscalex102 Marks, D.B.
, A.D. Marks, C.M. Smith. 2000. Basic Medical Biochemistry: A Clinical \line\tab
\up0 \expndtw-2\charscalex100 Approach. Terjemahan B.U. Pendit. Penerbit Buku K
edokteran EGC. Jakarta\ul0\nosupersub\cf1\f2\fs24 . \par\pard\ql \li1800\sb0\sl300\slmult0 \par\pard\ql\li1800\ri1625\sb20\sl-300\slmult0\tx2339\tx2339 \up0 \e
xpndtw0\charscalex103 \ul0\nosupersub\cf6\f7\fs22 Martin, T.W., Weisman, I.M., Z
eballos, R.J., and Stephenson, S.R. 1989. Exercise \line\tab \up0 \expndtw0\char
scalex110 and hypoxia increase sickling in venous blood from an exercising limb
in \line\tab \up0 \expndtw-3\charscalex100 individuals with sickle cell trait. A
m. J. Med. 87:48-56. \par\pard\ql \li1799\sb0\sl-306\slmult0 \par\pard\ql\li1799
\ri1625\sb10\sl-306\slmult0\tx2339\tx2339\tx2340 \up0 \expndtw0\charscalex100 Mo
diano, D., Luoni, G., Sirima, B.S., Simpore, J., Verra, F., Konate, A., Rastrell
i, E., \line\tab \up0 \expndtw0\charscalex102 Olivieri, A., Calissano, C., Pagan
otti, G.M., D'Urbano, L., Sanou, I., Sawadogo, \line\tab \up0 \expndtw0\charscal
ex100 A., Modiano, G., and Coluzzi, M. 2001. Haemoglobin C protects against clin
ical \line\tab \up0 \expndtw-3\charscalex100 Plasmodium falciparum malaria. Natu
re. 414:305-308. \par\pard\qj \li1800\sb0\sl-300\slmult0 \par\pard\qj\li1800\ri1
624\sb19\sl-300\slmult0\tx2339 \up0 \expndtw0\charscalex100 Murray, R.K., K.G.
Daryl, A.M. Peter and W.P. Victor. Harper\u8217?s Biochemistry. \line\t
ab \up0 \expndtw-3\charscalex100 Terjemahan A. Hartono. Penerbit Buku Kedokteran
EGC. Jakarta \par\pard\ql \li1800\sb0\sl-253\slmult0 \par\pard\ql\li1800\sb106\
sl-253\slmult0\tx6735 \up0 \expndtw0\charscalex103 Rank, B.H., Carlsson, J.,
and Hebbel, R.P. \tab \up0 \expndtw0\charscalex102 1985. Abnormal redox st
atus of \par\pard\ql \li2340\sb47\sl-253\slmult0 \up0 \expndtw-2\charscalex100
membrane-protein thiols in sickle erythrocytes. J. Clin. Invest. 75: 1531-1537 \
par\pard\ql \li1800\sb0\sl-300\slmult0 \par\pard\ql\li1800\ri1625\sb29\sl-300\sl
mult0\tx2340\tx2340\tx2339 \up0 \expndtw-1\charscalex100 Reed, M.B., Caruana, S.
R., Batchelor, A.H., Thompson, J.K., Crabb, B.S., and Cowman, \line\tab \up0 \ex
pndtw-3\charscalex100 A.F. \up0 \expndtw-2\charscalex100 2000. Targeted disr
uption of an erythrocyte binding antigen in \line\tab \up0 \expndtw0
\charscalex100 Plasmodium falciparum is associated with a switch toward
a sialic acid-\line\tab \up0 \expndtw-2\charscalex100 independent pathway of i
nvasion. Proc. Natl. Acad. Sci. USA. 97: 7509-7514. \par\pard\ql \li1800\sb0\sl306\slmult0 \par\pard\ql\li1800\ri1625\sb10\sl-306\slmult0\tx2340\tx2340\tx2340
\up0 \expndtw0\charscalex100 Ringelhann, B., Hathorn, M.K., Jilly, P., Grant, F.
, and Parniczky, G. 1976. A new look \line\tab \up0 \expndtw0\charscalex105 at t
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