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MITOKONDRIA Adnan (Dosen Biologi FMIPA Universitas Negeri Makassar)
A. PENDAHULUAN Energi yang langsung dikonsumsi oleh sel adalah energi dalam bentuk adenosin trifosfat atau ATP. Substansi yang kaya energi tersebut diubah menjadi adenosin difosfat atau ADP dan selanjutnya diubah menjadi adenosin monofosfat atau AMP. Peristiwa tersebut dike-nal dengan nama hidrolisis ATP. Rumus kimia ATP ditun-jukkan pada gambar 1.
Gambar 11.1. Rumus kimia ATP
Sel memiliki tiga jalur utama untuk menghasilkan energi utama dalam bentuk ATP, yaitu: 1. ATP dihasilkan di dalam sitosol selama glikolisis; 2. ATP dihasilkan di dalam kloroplas dengan memanfaatkan energi matahari; 3. ATP dihasilkan di dalam mitokondria melalui reaksi oksidasi substrat.
B. TEORI ENDOSIMBIOSIS Para pakar biologi sel sependapat bahwa mitokondria berasal dari bakteri aerobik yang mengadakan endosimbiosis dengan eukariot anaerobik. Endosimbiosis adalah jika
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organisme dari suatu species hidup didalam organisme dari species yang lain. Proses yang terjadi adalah sel-sel bakteri aerobik tertelan oleh sel eukariot anaerobik namun bakteri aerobik tidak mengalami pencernaan. Bakteri aerobik yang tertelan selanjutnya hidup di dalam sel eukariot. Eukariot mensuplai bakteri dengan proteksi dan komponen-komponen karbon, sedangkan bakteri mensuplai eukariot dengan energi ATP. Beberapa bukti yang mendukung teori tersebut tersebut adalah:(i) Mitokondria dapat menggandakan diri menyerupai pembelahan biner pada bakteri, (ii) DNA mitokondria menyerupai DNA prokariot, berupa molekul sirkular tunggal dan ribosom pada mitokondria dibuat dari sub unit dengan koofisien sedimentasi yang lebih menyerupai ribosom prokariot dibandingkan dengan eukariot, (iii) Ukuran mitokondria hampir sama dengan ukura bakteri. (iv) Mitokondria memiliki membran ganda, memiliki kemi-ripan dengam membran sejumlah prokariota, dan (v) urutan DNA pada mitokondria tertutup dan memiliki kemiripan dengan DNA proteobakteril (vi) Mitokondria memiliki kemiripan fisik dengan prokariota.
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Gambar 11.2. Teori endomembran http://www.biology.hawaii.edu/301/Lecture%20Powerpoints/Lecture%207%2 0Evolution%20and%20analyses%20of%20life%20histories.ppt
Mitokondria pertama kali diamati dan diisolasi dari sel pada tahun 1850 oleh Kollicker melalui pengamatannya pada jaringan otot lurik lurik serangga. serangga. Ia menemukan menemukan adanya adanya granulagranulagranula dengan struktur yang bebas dan tidak berhubungan secara langsung dengan struktur internal sel. Pada tahun 1890, 1890, Altmann mengidentifikasi granula-granula tersebut dan Ia berikan nama bioblast. Istilah tersebut diganti dengan mitokondria (Yunani: mito yang berarti benang dan chondrion yang berarti granula) sebab penampakan granula-granula tersebut menyerupai benang bila diamati dengan menggunakan mikroskop cahaya. Pada tahun 1900, Michaelis menunjukkan bahwa di dalam mitokondria berlangsung reaksi-reaksi oksidatif. Pada tahun 1911, Warburg menemukan bahwa mitokondria mengandung enzim-enzim yang mengkatalisis reaksi-reaksi oksidatif sel. sel. Pada tahun tahun 1911, Kingsbury mendu-kung bahwa mitokondria merupakan tempat spesifik untuk reaksi-reaksi oksidasi. Pada tahun 1930, Sir Hans Krebs menjelaskan beberapa reaksi siklus asam trikarboksilat atau daur Krebs. Dari tahun 1950, Lehninger, Green, Kennedy, dan Hogeboom dan lain-lain menunjukkan secara jelas reaksi-reaksi seperti oksidasi asam lemak, fosforilasi oksidatif serta sifat-sifat lain mitokondia (Sheeler dan Bianchii, 1983). C. STRUKTUR MITOKONDRIA Mitokondria dijumpai baik pada sel hewan maupun pada sel tumbuhan. Ukuran mitokondria kira-kira sama dengan ukuran rata-rata bakteri basil. Mitokondria hati secara umum agak memanjang dengan diameter kira-kira 0,5-1,0 µm dan panjang kira-kira 3 µm. Umumnya panjang mitokondria dapat mencapai 7 µm (Sheeler & Bianchi, 1983; Thorpe, 1984). Di dalam sel mitokondria terletak secara acak seperti pada hati atau tersusun teratur dengan pola-pola tertentu seperti pada sel sperma. Contoh yang paling umum adalah susunan yang teratur dari mitokondria diantara serabut-serabut di dalam otot lurik. Mitokondria umumnya ditemukan pada
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Untuk melaksanakan fungsinya, sangat tergantung pada persediaan ATP yang dihasilkan oleh mitokondria.
Gambar 11.3a. Mitokondria
Gambar 11.3b Susunan mitokondria pada sel otot lurik (Thorpe, 1984)
Gambar 11.3c. Susunan mitokondria pada ekor sel sperma (Thorpe, 1984)
Jumlah mitokondria per sel sangat bervariasi diantara berbagai tipe sel, mulai dari nol sampai ratusan ribu. Algae tak
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5
memiliki mitokondria rata-rata 800 per sel dan beberapa telur landak laut dan amoeba raksasa Chaos chaos mengandung 500.000 mitokondria per sel. Dalam beberapa hal, tampaknya terdapat hubungan antara jumlah mitokondria per sel dan keperluan metabolisme sel.
Gambar 11.4. 11.4. Struktur Mitokondria (Thorpe, 1984)
Mitokondria dibatasi oleh membran ganda, yaitu membran dalam dan membran luar. Setiap membran memiliki ciri khas sebagai unit membran. Membran dalam tidak berhubungan dengan membran luar. Membran dalam membagi organel menjadi dua bagian yaitu matriks dan ruang antar membran. Matriks berisi cairan menyerupai gel, sedangkan ruang antar membran berisi cairan yang encer. Membran dalam memiliki permukaan yang lebih luas dibandingkan dengan membran luar, karena membran dalam terlipat-lipat dan masuk ke dalam matriks membentuk tonjolan-tonjolan yang dinamakan krista. Dengan demikian, secara struktural terdapat perbedaan antara membran dalam dengan membran luar. Selain itu, membran dalam berbeda dengan membran luar dari segi permiabilitasnya. Membran luar permiabel terhadap berbagai substansi yang mempunyai berat molekul berkisar 5.000 dalton. Sebaliknya permiabilitas membran dalam terbatas, khususnya terhadap substansi-substansi dengan berat molekul berkisar 100-150 dalton (Sheeler & Bianchi, 1983). Struktur morfologi mitokondria yang paling bervariasi adalah krista. Dalam satu tipe sel, mereka pada umumnya uniform dan khas pada sel. Akan tetapi, susunan dari bentukbentuk yang berbeda terdapat dalam tipe-tipe sel yang berbeda.
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dan teratur, sedang pada krista yang berbentuk tubular memperlihatkan tubulus-tubulus yang terorientasi pada matriks. Pada beberapa mitokondria, susunan tubulusnya teratur, misalnya pada Amoeba Chaos chaos. Menurut Sheeler & Bianchi (1983), struktur mitokondria dapat dikelompokkan menjadi tiga, yaitu (i) krista susunannya menyerupai lembaran misalnya krista pada mitokondria sel hati, (ii) krista dengan susunan yang sangat rapat menyerupai tumpukan uang logam misalnya pada mitokondria sel ginjal, dan (iii) krista dengan susunan seperti jala yang dibentuk oleh saluran-saluran yang saling beranastomosis.
Gambar 11.5 Struktur krista mitokondria (Sheler dan Bianchii,1983)
D. KOMPOSISI KIMIA MITOKONDRIA Pada mitokondria utuh, air merupakan komponen utama yang dominan dan ditemukan di seluruh mitokondria kecuali dalam lapisan bilayer lipida. Air selain berperan dalam reaksireaksi kimia, juga berperan sebagai medium fisik dimana metabolit dapat berdifusi diantara sistim-sistim enzim. Komponen utama mitokondria adalah protein. Persentase protein yang sebenarnya berkaitan dengan jumlah membran dalam yang ada. Membran dalam terdiri atas protein, baik protein enzimatik maupun protein struktural. Pada beberapa mitokondria, membran dalam mengandung kira-kira 60% dari total protein organel. Berdasarkan distribusi enzim di dalam mitokondria hati tikus, telah terbukti bahwa membran dalam mengandung 21% dari total protein mitokondria dan membran luar 40%. Menurut perhitungan ini, kurang lebih 67% protein terdapat pada matriks dan biasanya ditemukan dalam ruang intraseluler.
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perifer membran atau protein intrinsik membran tertentu. Protein tidak terlarut biasanya menjadi bagian integral membran. Beberapa dari protein ini merupakan protein struktural serta beberapa protein enzim. Komposisi lipida mitokondria tergantung dari sumber mitokondrianya. Namun demikian, fosfolipida merupakan bentuk yang dominan. Umumnya fosfolipida terdiri dari ¾ dari total lipida (tabel 11.1). Tabel 11.1 Kandungan lipida mitokondria yang diisolasi dari berbagai organ hewan (dalam mg/g) (Thorpe, 1984)
Organ
Total Lipida
Lipida Netral
Kolest erol
Fosfolipida
Jantung manusia
400
-
-
335
Jantung sapi
320
18
4
283
Ginjal sapi
240
17
11
190
Hati sapi
180
16
4
145
-
-
-
159
Hati marmut
Fosfatidilamin dan fosfatidiletanolamin umumnya merupakan fosfatidil dalam jumlah yang besar pada mitokondria. Namun demikian, ditemukan kadar kardiolipin dan kolesterol dengan konsentrasi yang rendah (tabel 11.2) Tabel 11.2 Komposisi fosfolipida mitokondria yang diisolasi dari berbagai sumber (% total fosfolipida) (Thorpe, 1984)
Sumber
Fosfatidil kolin
Fosfatidil etanolami n
Kardi olipin
Fosfati dil inositol
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Distribusi fosfolipida dalam membran luar dan membran dalam mitokondria ditunjukkan pada tabel 11.3. Tabel 11.3 Komposisi fosfolipida membran dalam dan membran luar mitokondria hati (% fosfolipida dalam setiap fraksi) (Thorpe, 1984)
Marmut
Sumber
Tikus
M. dalam
M. luar
M. dalam
M. luar
Fosfatidil kolin
44,50
55,20
41,00
39,00
Fosfatidil etanolamin
25,30
27,70
35,00
31,00
Kardiolipin Kardiolip in
21,50
3,20
21,00
3,00
Fosfatidil inositol
4,20
13,50
-
-
Fosfatidil serin
-
-
2,00
17,00
Perbedaan distribusi lipida memiliki arti penting, baik dari segi struktural maupun fungsional. Namun secara detail belum jelas. Sejumlah molekul organik sederhana yang berbeda berasosiasi dengan membran mitokondria. Beberapa dari molekul ini adalah molekul redoks yang ikut serta dalam transpor elektron. Ubiquinon (koenzim Q), flavin (FMN dan FAD), dan nukleotida piridin (NAD +) secara normal terikat membran, dan kadang-kadang berasosiasi pada hampir sebahagian besar membran dalam. E. KOMPARTEMEN ENZIM Kurang lebih 100 enzim telah diidentifikasi berhubungan dengan mitokondria. Kira-kira 37% dari enzim-enzim tersebut
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Gambar 11.6. Komprtemen enzim pada mitokondria (Thorpe,1984)
Matriks mengandung sekumpulan enzim yang merupakan mediator reaksi siklus asam trikarboksilat (TCA) dan berkaitan dengan sintesis protein dan asam nukleat. Semua enzim-enzim TCA bebas di dalam matriks kecuali suksinat dehidrogenase, yang merupakan satu komponen membran dalam. Jadi untuk piruvat, dioksidasi sempurna menjadi CO 2 dan H2O di dalam matriks. Metabolisme suksinat harus mengadakan kontak dengan membran dalam sebelum
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Gambar 11.7. Siklus TCA. Suksinat dehidrogenase merupakan satusatunya enzim yang terikat membran (Thorpe, 1984)
F. GLIKOLISIS Respirasi seluler merupakan rangkaian peristiwa yang berlangsung melalui pemecahan glukosa menjadi asam piruvat, perubahan asam piruvat menjadi asetil KoA, daur krebs dan rantai pernapasan. Walaupun glikolisis berlangsung di dalam sitoplasma, namun sebagai rangkaian dari proses respirasi seluler, maka pada uraian berikut ini juga akan dibahas mengenai glikolisis. Glikolisis adalah proses penguraian molekul heksosa yang memiliki enam atom karbon dan berlangsung secara enzimatis untuk menghasilkan dua molekul asam piruvat yang
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11
energi dalam bentuk ATP. Fase persiapan terdiri atas lima tahapan reaksi. Heksosa lain seperti seperti D-fruktosa, D-fruktosa, D-Galaktosa, D-Galaktosa, dan D-mannosa dapat masuk ke dalam fase persiapan glikolisis setelah mengalami fosfo-rilasi. Fase produksi energi " berlangsung melalui lima tahapan reaksi berikutnya. Dalam peristiwa ini dihasilkan 4 molekul ATP. Pada tahap awal glikolisis, glukosa diubah menjadi fruktosa 1,6 bifosfat dengan memanfaatkan dua molekul ATP. Fruktosa 1,6 bifosfat dipecah menjadi 2 molekul senyawa 3 C yaitu dihidroksi aseton fosfat dan gliseraldehida 3 fosfat yang keduanya merupakan isomer gliseraldehida 3 fosfat. Selanjutnya mengalami reaksi dengan Pi kemudian diikuti dengan reaksi reduksi pembentukan NADP dari NAD dan terbentuk asam 1,3 difosfogliserat.
Glukosa 6 Fosfat
Fruktosa 6 Fosfat
Fruktosa 1,6 difosfat
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asam piruvat. Pada perubahan asam 1,3 difosfogliserat menjadi 3 fosfogliserat dan dari fosfoenol piruvat menajdi asam piruvat dirangkaikan dengan pembentukan ATP dari ADP dan Pi yang dilepaskan. Seluruh reaksi perubahan glukosa sehingga terbentuk asam piruvat melibatkan berbagai enzim sesuai substrat yang bereaksi. Seluruh rangkaian respirasi menghasilkan 2 molekul ATP dan 2 NADPH. Selama berlangsungnya glikolisis , terdapat tiga jenis transformasi kimia yang berbeda, yaitu: − Pemecahan kerangka karbon glukosa menghasilkan asam piruvat. − Fosforilasi ADP menjadi ATP oleh senyawa fosfat berenergi tinggi yang dibentuk selama glikolisis. − Pemindahan atom hidrogen atau elektron. Menurut Sheeler dan Bianchii (1983), ada empat ciri utama gliklisis, yaitu: −
−
Gula pertama menglami dua kali fosforilasi. Pada gula seperti glukosa, fruktosa dan mannosa membutuhkan dua molekul ATP per mol monosakarida. Sedangkan gula yang diturunkan dari glikogen atau pati, hanya membutuhkan satu mol ATP permol permol glukosa glukosa equivalen. equivalen. Jadi fosfat anorganik dibutuhkan selama fosforilasi polisakarida. Gula difosfat berkarbon enam dipecah oleh enzim
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13
Gambar 11.9. Glikolisis tahap kedua Tahap-tahap reaksi kimia glikolisis secara kseluruhan ditunjukkan sebagai berikut: 1. Reaksi pemindahan pemindahan fosfat. fosfat. Enzim kinase memindahkan memindahkan fosfat fosfat dari ATP suatu akseptor. Enzim heksokinase pada umumnya lebih spesifik untuk memindahkan fosfat ke glukosa. gl ukosa.
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3. Reaksi pemindahan fosfat. Reaksi ini dibantu oleh enzim fosfofruktokinase.
4. Pemecahan karbohidrat enam karbon menjadi 3 carbon. Reaksi ini dibantu oleh enzim aldolase.
5. Perubahan DHAP menjadi PGAL dengan bantuan enzim triosa fosfat isomerase.
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15
8. Perubahan 3-fosfogliserat menjadi 2-fosfogliserat dengan bantuan enzim fosfogliserat mutase.
9. Hidrolisis 2-fosfogliserat menjadi fosfoenolpruvat dengan bantuan enzim enolase.
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dikonversi menjadi etanol atau asam laktat dalam kondisi anaerb.
Gambar 11.10. 11.10. Kemungkinan Proses Lanjut Asam Piruvat
1. Fermentasi Etanol Fermentasi etanol dari asam piruvat berlangsung dalam keadaan anaerob. anaerob. Proses ini dapat berlangsung berlangsung pada ragi dan beberapa beberapa mikroorganisme lainnya.
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17
Gambar 11.13. 11.13. Koenzim NAD
Hasil akhir konversi gula menjadi etanol disebut fermentasi alkohol. alkohol. Hasil bersih proses anaerob anaerob ini adalah: adalah: + Glukosa + 2Pi + 2ADP + 2H 2 etanol +2CO2 +2ATP+2H2O Hal penting untuk diperhatikan adalah bahwa NAD + dan
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Gambar 11.15. Fermentasi laktat
Reaksi keseluruhan pada konversi glukosa menjadi laktat adalah: Glukosa + 2Pi + 2AP 2 laktat + 2ATP+2H2O Asam laktat dan asam piruvat, di dalam sel dapat
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Tabel 11.1. Sub unit kompleks piruvat dehidrogenase
Pyruvate Dehydrogenase SCoA HSCo A
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21
Thiamine pyrophosphate (TPP TPP)) adalah turunan dari tiamin atau vitamin B1..
S
CH2 CH2
S
CH
lipoic acid
O
lysine
NH
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23
dalam daur krebs adalah CO 2, dan air. Selain itu dihasilkan sejumlah komponen-komponen tertentu seperti NADH yang berpartisipasi di dalam rantai respirasi atau rantai transpor elektron dan secara khusus berhubungan dengan membran dalam mitokondria. Hasil dari reaksi-reaksi yang berlangsung selama rantai respirasi adalah reduksi O 2 untuk membentuk H2O. Selain itu selama berlangsungnya rantai respirasi juga berlangsung fosforilasi oksidatif yang mengubah ADP menjadi
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Ringkasan reaksi-reaksi yang terjadi di dalam daur Krebs ditunjukkan pada gambar 11.9. Daur Krebs memiliki keterkaitan yang sangat erat dengan rantai respirasi. Di dalam rantai respirasi berlangsung reaksi oksidasi reduksi yang menghantarkan elektron dan H + ke O2 untuk membentuk air. Seiring dengan berlangsungnya rantai respirasi, juga berlangsung fosforilasi oksidatif yang mengubah ADP menjadi ATP (gambar 11.11).
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NADH + H+ + ADP + Pi + ½ O2
NAD+H2O + ATP
FADH2 atau FMNH2 selanjutnya memindahkan 2 elektron dan 2 H+ ke suatu enzim yang mengandung besi (Fe) yang terikat pada gugus SH. Hal ini mengakibatkan enzim tersebut tersebut +++ tereduksi dan menyebabkan Fe (ferro) teroksidasi menjadi ++ Fe (feri). Selanjutnya dari enzim ini memindahkan 2 elektron dan 2 H+ ke ubiquinon (UQ).
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elektron dibedakan atas sitokrom b, sitokrom c, dan sitokrom
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Fosforilasi
Fosforilasi
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Mekanisme transkripsi dan translasi pada mitokondria