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== Referensi ==
== Referensi ==
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<ref name=Little2008>{{cite journal |author=Little AEF, Currie CR. |title=Black yeast symbionts compromise the efficiency of antibiotic defenses in fungus-growing ants |journal=Ecology |year=2008 |volume=89 |issue=5 |pages=1216–1222 |doi=10.1890/07-0815.1 |pmid=18543616}}</ref>

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<ref name="Magliani 2005">{{cite journal |author=Magliani 1, Conti S, Frazzi R, Ravanetti L, Maffei DL, Polonelli L. |title=Protective antifungal yeast killer toxin-like antibodies |journal=Current Molecular Medicine |year=2006 |volume=5 |issue=4 |pages=443–52 |pmid=15978000 |doi=10.2174/1566524054022558}}</ref>

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<ref name=McBryde>{{cite journal |author=McBryde C, Gardner JM, de Barros Lopes M, Jiranek V. |year=2006 |title=Generation of novel wine yeast strains by adaptive evolution |journal=American Journal of Enology and Viticulture |volume=57 |issue=4 |pages=423–430}}</ref>

<ref name="McFarland 2010">{{cite journal |author=McFarland LV. |title=Systematic review and meta-analysis of ''Saccharomyces boulardii'' in adult patients |journal=World Journal of Gastroenterology |year=2010 |volume=16 |issue=18 |pages=2202–22 |pmid=20458757 |pmc=2868213 |doi=10.3748/wjg.v16.i18.2202}} {{open access}}</ref>

<ref name="MooreLandecker1996p533-534">Moore-Landecker, pp. 533–534.</ref>

<ref name=NASA>{{cite web |url=http://science.nasa.gov/newhome/headlines/msad16mar99_1b.htm |title=Planets in a bottle: more about yeast |work=Science@NASA |author=Phillips T |accessdate=15 January 2012}}</ref>

<ref name=Nielsen 2012">{{cite journal |author=Nilesen J. |title=Production of biopharmaceutical proteins by yeast. Advances through metabolic engineering |journal=Bioengineered |year=2012 |volume=4 |issue=4 |pages=207–211 |doi=10.4161/bioe.22856}} {{open access}}</ref>

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<ref name="Siddiqui 2012">{{cite journal |author=Siddiqui MS, Thodey K, Trenchard I, Smolke CD. |title=Advancing secondary metabolite biosynthesis in yeast with synthetic biology tools |journal=FEMS Yeast Research |year=2012 |volume=12 |issue=2 |pages=144–170 |doi=10.1111/j.1567-1364.2011.00774.x}} {{open access}}</ref>

<ref name=Slavikova2003>{{cite journal |author=Sláviková E, Vadkertiová R. |title=The diversity of yeasts in the agricultural soil |journal=Journal of Basic Microbiology |volume=43 |issue=5 |pages=430–436 |year=2003 |pmid=12964187 |doi=10.1002/jobm.200310277}}</ref>

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<ref name="Soares 2012">{{cite journal |author=Soares EV, Soares HMVM. |title=Bioremediation of industrial effluents containing heavy metals using brewing cells of ''Saccharomyces cerevisiae'' as a green technology: A review |journal=Environmental Science and Pollution Research |year=2012 |volume=19 |issue=4 |pages=1066–83 |doi=10.1007/s11356-011-0671-5 |pmid=22139299}}</ref>

<ref name=Suh2005>{{cite journal |author=Suh SO, McHugh JV, Pollock DD, Blackwell M. |title=The beetle gut: a hyperdiverse source of novel yeasts |journal=Mycological Research |volume=109 |issue=3 |pages=261–265 |year=2005 |pmid=15912941 |pmc=2943959 |doi=10.1017/S0953756205002388}}</ref>

<ref name=Teoh2004>{{cite journal |author=Teoh AL, Heard G, Cox J. |year=2004 |title=Yeast ecology of Kombucha fermentation |journal=International Journal of Food Microbiology |volume=95 |issue=2 |pages=119–126 |doi=10.1016/j.ijfoodmicro.2003.12.020 |pmid=15282124}}</ref>

<ref name="Thaler 2014">{{cite book |author=Thaler M, Safferstein D. |title=A Curious Harvest: The Practical Art of Cooking Everything |url=http://books.google.com/books?id=lD_0AwAAQBAJ&pg=PA129 |year=2014 |publisher=Quarry Books |isbn=978-1-59253-928-4 |page=129}}</ref>

<ref name="Tong 2006">{{cite book |editor-last=Xiao W. (ed.) |title=Yeast Protocols |url=http://books.google.com/books?id=sC0Am-HWeeEC&pg=PA171 |author=Tong AHY, Boone C. |chapter=Synthetic genetic array analysis in ''Saccharomyces cerevisiae'' |year=2006 |publisher=Springer Science & Business Media |isbn=978-1-59259-958-5 |pages=171–191}}</ref>

<ref name="urlLe Comité des Fabricants de levure&nbsp;— COFALEC&nbsp;— représente les fabricants">{{cite web |url=http://www.cofalec.com/contentO.aspx?progres |title=Le Comité des Fabricants de levure |publisher=COFALEC |accessdate=21 February 2010}}</ref>

<ref name="urlUMMC">{{cite web |title=Brewer's yeast |url=http://www.umm.edu/altmed/articles/brewers-yeast-000288.htm |publisher=University of Maryland Medical Center |accessdate=15 January 2012}}</ref>

<ref name=Vanderhaegen2003>{{cite journal |author=Vanderhaegen B, Neven H, Cogne S, Vertrepin KJ, Derdelinckx C, Verachtert H. |year=2003 |title=Bioflavoring and Beer Refermentation |journal=Applied Microbiology and Biotechnology |volume=62 |issue=2–3 |pages=140–150 |doi=10.1007/s00253-003-1340-5|pmid=12759790 |last2=Neven |last3=Coghe |last4=Verstrepen |last5=Derdelinckx |last6=Verachtert }}</ref>

<ref name=VanderWalt1984>{{cite journal |author=Van der Walt JP. |year=1984 |title=The Yeasts: A Taxonomic Study |edition=3rd |journal=Elsevier Science, Amsterdam |pages=146–150}}</ref>

<ref name=Walker>{{cite journal |author=Walker K, Skelton H, Smith K. |title=Cutaneous lesions showing giant yeast forms of ''Blastomyces dermatitidis'' |journal=Journal of Cutaneous Pathology |volume=29 |issue=10 |pages=616–618 |year=2002 |pmid=12453301 |doi=10.1034/j.1600-0560.2002.291009.x}}</ref>

<ref name=Williams1996>{{cite journal |author=Williams N. |year=1996 |title=Genome Projects: Yeast genome sequence ferments new research |journal=Science |volume=272 |issue=5261 |page=481 |doi=10.1126/science.272.5261.481 |pmid=8614793|bibcode=1996Sci...272..481W }}</ref>

<ref name=Wood2002>{{cite journal |vauthors=Wood V, Gwilliam R, Rajandream MA, etal |title=The genome sequence of ''Schizosaccharomyces pombe'' |journal=[[Nature (journal)|Nature]] |volume=415 |issue=6874 |pages=871–880 |year=2002 |pmid=11859360 |doi=10.1038/nature724}}</ref>

<ref name="YeastRef1">{{cite book |author=Kurtzman CP, Fell JW. |year=2006 |url=http://www.ars.usda.gov/research/publications/publications.htm?SEQ_NO_115=176765 |chapter=Yeast Systematics and Phylogeny—Implications of Molecular Identification Methods for Studies in Ecology |title=Biodiversity and Ecophysiology of Yeasts, The Yeast Handbook, |publisher=Springer}}</ref>

<ref name=Yakobson2010>{{cite journal |author=Yakobson CM. |year=2010 |title=Pure culture fermentation characteristics of ''Brettanomyces'' yeast species and their use in the brewing industry |url=http://www.brettanomycesproject.com/dissertation/}}</ref>

<ref name="YeastRef3">{{cite web |url=http://automation.tkk.fi/Research/BioprocessAutomation |title=Bioprocess automation |publisher=Helsinki University of Technology |year=2007 |accessdate=15 January 2012}}</ref>

<ref name="YeastRef2">{{cite web |url=http://www.yeastgenome.org/VL-what_are_yeast.html |title=What are yeasts? |work=Yeast Virtual Library |date=13 September 2009 | accessdate=28 November 2009}}</ref>

<ref name="YeastRef4">{{cite web|url=http://genomicsgtl.energy.gov/biofuels/ethanolproduction.shtml |title=Fuel Ethanol Production: GSP Systems Biology Research |publisher=U.S. Department of Energy Office of Science |work=Genomic Science Program |accessdate=28 November 2009 |deadurl=yes |archiveurl=https://web.archive.org/20090603231351/http://genomicsgtl.energy.gov/biofuels/ethanolproduction.shtml |archivedate=3 June 2009 }}</ref>

<ref name="YeastRef8">{{cite web |url=http://www.cofepow.org.uk/pages/asia_haruku2.htm |title=South East Asia Under Japanese Occupation&nbsp;– Harukoe (Haruku) |author=Lee JG (ed.) |work=Children (& Families) of the Far East Prisoners of War |accessdate=28 November 2009}}</ref>

<ref name="YeastRef9">{{Wayback |date=20060925065545 |url=http://helios.bto.ed.ac.uk/bto/microbes/yeast.htm |title="The Microbial World: Yeasts and yeast-like fungi" }}. ''Institute of Cell and Molecular Biology''. Retrieved 24 December 2006.</ref>

<ref name=YeastRef11>{{cite web |author=Huxley A | url=http://aleph0.clarku.edu/huxley/CE8/Yeast.html |title=Discourses: Biological & Geological (volume VIII) : Yeast |year=1871 |work=Collected Essays |accessdate=28 November 2009}}</ref>

<ref name=YeastRef14>{{Wayback |date=20060921160928 |url=http://www.gwrdc.com.au/downloads/0506docs/researcher.pdf |title=Research enables yeast suppliers to expand options }}. Retrieved 10 January 2007.</ref>

<ref name=Yeong2005>{{cite journal |author=Yeong FM. |title=Severing all ties between mother and daughter: cell separation in budding yeast |journal=Molecular Microbiology |volume=55 |issue=5 |pages=1325–1331 |year=2005 |pmid=15720543 |doi=10.1111/j.1365-2958.2005.04507.x}}</ref>

<ref name=YeastRef15>{{cite web |url=http://www.winepros.org/wine101/vincyc-bret.htm |title=''Brettanomyces'' (Dekkera) |author=Lamar J |work=Vincyclopedia |accessdate=28 November 2009}}</ref>

<ref name="Zinjard 2014">{{cite journal |author=Zinjarde S, Apte M, Mohite P, Kumar AR. |title=''Yarrowia lipolytica'' and pollutants: Interactions and applications |journal=Biotechnology Advances |year=2014 |volume=32 |issue=5 |pages=920–933 |doi=10.1016/j.biotechadv.2014.04.008 |pmid=24780156}}</ref>

}}

=== Cited texts ===
* {{cite book |author=Alexopoulos CJ, Mims CW, Blackwell M. |title=Introductory Mycology |publisher=Wiley |location=New York, New York |year=1996 |isbn=0-471-52229-5}}
* {{cite book |author=Kirk PM, Cannon PF, Minter DW, Stalpers JA. |title=Dictionary of the Fungi |edition=10th |publisher=CAB International |location=Wallingford, UK |year=2008 |isbn=0-85199-826-7}}
* {{cite book |editor-last=Kurtzman CP, Fell JW, Boekhout T. |title=The Yeasts: A Taxonomic Study |volume=1 |edition=5th |year=2011 |publisher=Elsevier |location=Amsterdam, etc. |isbn=978-0-12-384708-9}}
* {{cite book |author=Moore-Landecker E. |title=Fundamentals of the Fungi |publisher=Prentice Hall |location=Englewood Cliffs, New Jersey |year=1996 |isbn=0-13-376864-3}}
*{{cite book |author=Priest FG, Stewart GG. |title=Handbook of Brewing |edition=2nd |url=http://books.google.com/books?id=TIYbNdrIsPEC&pg=PA691 |year=2006 |publisher=CRC Press |isbn=978-1-4200-1517-1 |page=691}}
=== Catatan ===
=== Catatan ===
* {{cite book |author=Alexopoulos CJ, Mims CW, Blackwell M. |title=Introductory Mycology |publisher=Wiley |location=New York, New York |year=1996 |isbn=0-471-52229-5}}
* {{cite book |author=Alexopoulos CJ, Mims CW, Blackwell M. |title=Introductory Mycology |publisher=Wiley |location=New York, New York |year=1996 |isbn=0-471-52229-5}}

Revisi per 10 November 2015 15.39

Diagram sel khamir

Khamir adalah mikroorganisme eukariot yang diklasifikasikan dalam kingdom Fungi, dengan 1.500 species yang telah dapat dideskripsikan[1] (diperkirakan 1% dari seluruh spesies fungi).[2] Khamir merupakan mikroorganisme uniseluler, meskipun beberapa spesies dapat menjadi multiseluler melalui pembentukan benang dari sel-sel budding tersambung yang dikenal sebagai hifa semu(pseudohyphae), seperti yang terlihat pada sebagian besar kapang.[3] Ukuran kapang bervariasi tergantung spesies, umumnya memiliki diameter 3–4 µm,namun beberapa jenis khamir dapat mencapai ukuran lebih 40 µm.[4] Sebagian besar khamir bereproduksi secara aseksual dengan mitosis, dan dengan pembelahan sel asimetris yang disebut budding.

Khamir yang paling umum digunakan adalah Saccharomyces cerevisiae, yang dimanfaatkan untuk produksi anggur, roti, tape, dan bir sejak ribuan tahun yang silam dalam bentuk ragi. Saccharomyces cerevisiae dapat mengkonversi karbohidrat menjadi karbon dioksida dan alkohol melalui proses fermentasi, karbon dioksida digunakan dalam proses pembuatan roti (baking) dan alkohol dalam minuman beralkohol.[5] Saccharomyces cerevisiae juga merupakan organisme model penting dalam penelitian biologi sel modern, dan juga salah satu mikroorganisme eukariot yang paling sering diteliti secara menyeluruh. Peneliti menggunakannya untuk mendapatkan informasi mengenai biologi sel eukariot dan terutama biologi manusia.[6] Spesies khamir lainnya seperti Candida albicans adalah patogen oportunistik dan dapat menyebabkan infeksi pada manusia (kandidiasis) . Khamir juga dapat digunakan untuk menghasilkan listrik dalam microbial fuel cell,[7] dan memproduksi etanol untuk industri biofuel.

Khamir tidak membentuk kelompok taksonomi atau filogeni tunggal. Istilah "khamir" atau “ragi” sering digunakan sebagai sinonim dari Saccharomyces cerevisiae,[8] namun keragaman filogeni dari khamir dipisahkan dalam 2 filum terpisah: Ascomycota dan Basidiomycota. Khamir yang reproduksinya dengan budding ("khamir sejati") diklasifikasikan dalam ordo Saccharomycetales.[9]

Sejarah

Khamir mungkin merupakan salah satu organisme yang dipelihara paling awal. Manusia menggunakan khamir untuk fermentasi dan pembuatan roti. Arkeologis menemukan batu penggiling dan ruang pembuatan untuk roti yang dikembangkan dengan khamir juga gambar bakery dan brewery berumur 4000 tahun .[10] Pada tahun 1680, Anton van Leeuwenhoek pertama kali mengamati khamir dengan mikroskop, namun pada saat itu khamir tidak dianggap sebagai organisme yang hidup, namun hanya sebagai struktur berbentuk globula.[11] Pada tahun 1857, mikrobiologis Perancis, Louis Pasteur membuktikan dalam tulisannya yang berjudul "Mémoire sur la fermentation alcoolique" bahwa fermentasi alkohol dilakukan oleh khamir yang hidup dan bukan merupakan katalis kimiawi.[10][12] Pasteur menunjukkan bahwa dengan memberikan gelembung oksigen ke dalam larutan berisi khamir, pertumbuhan sel dapat meningkat, namun proses fermentasi terhambat, pengamatan tersebut disebut Pasteur effect.

Pada akhir abad ke-18, dua galur khamir digunakan dalam pembuatan bir diidentifikasi: Saccharomyces cerevisiae, yang disebut ‘’top-fermenting yeast’’, dan Saccharomyces carlsbergensis, ‘’bottom-fermenting yeast’’. S. cerevisiae telah dijual secara komersial oleh orang-orang Belanda untuk pembuatan roti sejak 1780; sekitar tahun 1800 orang-orang Jerman juga mulai memproduksi khamir dalam bentuk krim. Pada tahun 1825, sebuah metode telah dikembangkan untuk menghilangkan cairan sehingga khamir dapat dijual dalam bentuk blok padat.[13] Produksi blok khamir dalam skala industry diperkuat dengan diperkenalkannya ‘’filter press’’ pada tahun 1867. Pada tahun 1872, Baron Max de Springer mengembangkan proses produksi khamir berbentuk granula yang digunakan sampai perang dunia I.[14] Di Amerika Serikat, khamir yang secara alami terdapat di udara digunakan secara ekslusif sampai khamir komersial dipasarkan di Philadelphia, dimana Charles L. Fleischmann memamerkan produk, prosedur pemakaian, dan roti yang dihasilkan.[15]

Nutrisi dan pertumbuhan

Khamir adalah chemoorganotroph karena menggunakan senyawa organik sebagai sumber energi dan tidak membutuhkan cahaya matahari untuk pertumbuhannya. Sebagian besar karbon didapat dari gula heksosa seperti glukosa dan fruktosa, atau disakarida seperti sukrosa dan maltosa. Beberapa spesies dapat memetabolisme gula pentosa seperti ribosa,[16] alkohol, dan asam organik. Spesies khamir ada yang membutuhkan oksigen untuk respirasi seluler aerobik (aerob obligat) atau anaerobik, namun juga dapat menghasilkan energi secara aerobik (anaerob fakultatif). Tidak seperti bakteri, belum ada spesies khamir yang hanya dapat tumbuh secara anaerob (anaerob obligat). Khamir tumbuh dengan baik pada lingkungan pH netral atau sedikit asam. Suhu optimal pertumbuhan khamir bervariasi antar spesies . Sebagai contoh, Leucosporidium frigidum dapat bertumbuh pada −2 hingga 20 °C (28 hingga 68 °F), Saccharomyces telluris pada 5 hingga 35 °C (41 hingga 95 °F), dan Candida slooffi pada 28 hingga 45 °C (82 hingga 113 °F).[17] Sel dapat tetap bertahan hidup saat dibekukan dalam kondisi tertentu, namun dengan daya hidup yang menurun seiring waktu.

Umumnya, khamir ditumbuhkan di laboratorium pada media pertumbuhan padat maupun cair (broth). Media yang umum digunakan untuk menumbuhkan khamir adalah potato dextrose agar atau potato dextrose broth, Wallerstein Laboratories nutrient agar, yeast peptone dextrose agar, dan yeast mould agar atau broth. Pembuat minuman alkohol dalam skala rumahan umumnya menggunakan ekstrak malt dan agar sebagai media pertumbuhan padar. Antibiotik cycloheximide terkadang ditambahkan pada media pertumbuhan khamir untuk menghambat pertumbuhan khamir Saccharomyces dan menyeleksi spesies khamir liar/alami.

Lihat pula

Referensi

  1. ^ Kesalahan pengutipan: Tag <ref> tidak sah; tidak ditemukan teks untuk ref bernama YeastRef1
  2. ^ Kesalahan pengutipan: Tag <ref> tidak sah; tidak ditemukan teks untuk ref bernama Kurtzman2
  3. ^ Kesalahan pengutipan: Tag <ref> tidak sah; tidak ditemukan teks untuk ref bernama Kurtzman1
  4. ^ Kesalahan pengutipan: Tag <ref> tidak sah; tidak ditemukan teks untuk ref bernama Walker
  5. ^ Kesalahan pengutipan: Tag <ref> tidak sah; tidak ditemukan teks untuk ref bernama Legras2007
  6. ^ Kesalahan pengutipan: Tag <ref> tidak sah; tidak ditemukan teks untuk ref bernama Ostergaard
  7. ^ Kesalahan pengutipan: Tag <ref> tidak sah; tidak ditemukan teks untuk ref bernama YeastRef3
  8. ^ Kesalahan pengutipan: Tag <ref> tidak sah; tidak ditemukan teks untuk ref bernama Kurtzman1994
  9. ^ Kesalahan pengutipan: Tag <ref> tidak sah; tidak ditemukan teks untuk ref bernama YeastRef2
  10. ^ a b Kesalahan pengutipan: Tag <ref> tidak sah; tidak ditemukan teks untuk ref bernama NASA
  11. ^ Kesalahan pengutipan: Tag <ref> tidak sah; tidak ditemukan teks untuk ref bernama YeastRef11
  12. ^ Kesalahan pengutipan: Tag <ref> tidak sah; tidak ditemukan teks untuk ref bernama Barnett2003
  13. ^ Kesalahan pengutipan: Tag <ref> tidak sah; tidak ditemukan teks untuk ref bernama Klieger2004
  14. ^ Kesalahan pengutipan: Tag <ref> tidak sah; tidak ditemukan teks untuk ref bernama urlLe Comité des Fabricants de levure — COFALEC — représente les fabricants
  15. ^ Kesalahan pengutipan: Tag <ref> tidak sah; tidak ditemukan teks untuk ref bernama Snodgrass2004
  16. ^ Kesalahan pengutipan: Tag <ref> tidak sah; tidak ditemukan teks untuk ref bernama Barnett1975
  17. ^ Kesalahan pengutipan: Tag <ref> tidak sah; tidak ditemukan teks untuk ref bernama Arthur1976


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Cited texts

  • Alexopoulos CJ, Mims CW, Blackwell M. (1996). Introductory Mycology. New York, New York: Wiley. ISBN 0-471-52229-5. 
  • Kirk PM, Cannon PF, Minter DW, Stalpers JA. (2008). Dictionary of the Fungi (edisi ke-10th). Wallingford, UK: CAB International. ISBN 0-85199-826-7. 
  • Kurtzman CP, Fell JW, Boekhout T., ed. (2011). The Yeasts: A Taxonomic Study. 1 (edisi ke-5th). Amsterdam, etc.: Elsevier. ISBN 978-0-12-384708-9. 
  • Moore-Landecker E. (1996). Fundamentals of the Fungi. Englewood Cliffs, New Jersey: Prentice Hall. ISBN 0-13-376864-3. 
  • Priest FG, Stewart GG. (2006). Handbook of Brewing (edisi ke-2nd). CRC Press. hlm. 691. ISBN 978-1-4200-1517-1. 

Catatan

  • Alexopoulos CJ, Mims CW, Blackwell M. (1996). Introductory Mycology. New York, New York: Wiley. ISBN 0-471-52229-5. 
  • Kirk PM, Cannon PF, Minter DW, Stalpers JA. (2008). Dictionary of the Fungi (edisi ke-10th). Wallingford, UK: CAB International. ISBN 0-85199-826-7. 
  • Kurtzman CP, Fell JW, Boekhout T., ed. (2011). The Yeasts: A Taxonomic Study. 1 (edisi ke-5th). Amsterdam, etc.: Elsevier. ISBN 978-0-12-384708-9. 
  • Moore-Landecker E. (1996). Fundamentals of the Fungi. Englewood Cliffs, New Jersey: Prentice Hall. ISBN 0-13-376864-3. 
  • Priest FG, Stewart GG. (2006). Handbook of Brewing (edisi ke-2nd). CRC Press. hlm. 691. ISBN 978-1-4200-1517-1. 

Pranala luar


Ikon rintisan

Artikel bertopik Fungi ini adalah sebuah rintisan. Anda dapat membantu Wikipedia dengan mengembangkannya.

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  3. ^ Amendola J, Rees N. (2002). Understanding Baking: The Art and Science of Baking. John Wiley and Sons. hlm. 36. ISBN 978-0-471-40546-7. 
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