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Listeria Monocytogenes Soylarının Genetik Ve Virülens Farklılıkları

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Listeria Monocytogenes Soylarının Genetik Ve Virülens Farklılıkları Vet Hekim Der Derg 89(1): 97-107,2018 Çağrılı Makale / Invited Paper Listeria monocytogenes soylarının genetik ve virülens farklılıkları Nurcay KOCAMAN*, Belgin SARIMEHMETOĞLU** Öz: Listeria monocytogenes insanlarda ve Giriş hayvanlarda septisemi, menenjit, meningoensefalit, L. monocytogenes, gıda kaynaklı hastalık düşük gibi ciddi invazif hastalıklara neden olabilen oluşturan etiyolojik bir ajandır (29, 39, 40, 49). Gıda gıda kaynaklı bir patojendir. Epidemiyolojik zincirinde yüksek tuz konsantrasyonu, ekstrem pH ve araştırmalarda, gıda işletmelerinden kontaminasyon sıcaklık gibi koşullarda hayatta kalabilme özelliğine kaynağının takibinde ve farklı türler arasındaki sahiptir (1, 15, 22, 27). ilişkinin evriminin belirlenmesinde L. monocytogenes Kısa çubuk görünümünde, tek veya kısa zincir türlerinin alt tiplendirmesi çok önemlidir. Bu şeklinde, 0,4 - 0,5 x 1-2 µm boyutlarında, paralel derlemede L. monocytogenes’in soyları ile soyları kenarlı ve küt uçlu, Gram pozitif bir bakteri olan L. arasındaki genetik ve virülens farklılıklarından monocytogenes; L. grayi, L. innocua, L. ivanovii, L. bahsedilmiştir. welshimeri ve L. seeligeri ile birlikte Bacilli sınıfı, Anahtar sözcükler: Epidemiyoloji, Listeria Bacillales takımı, Listeriaceae familyasında yer monocytogenes, soy, virülens almaktadır (30). Son zamanlarda, geleneksel fenotipik Genetic and virulence differences of Listeria metodlar ve genom dizilimi kullanılarak yapılan monocytogenes strains araştırmalarda, Listeria soyunun bilinen 6 türünden Abstract: Listeria monocytogenes is a foodborne farklı olduğu belirtilen yeni türler bildirilmiştir. pathogen capable of causing serious invasive Graves ve ark. (16) Listeria marthii; Leclercq ve ark. disease, including abortion, septicemia, meningitis, (26) Listeria rocourtiae; Bertsch ve ark. (2) Listeria and meningoencephalitis in humans and animals. fleischmannii; Den Bakker ve ark. (11) Listeria Subtyping of L. monocytogenes strains can prove to floridensis, Listeria aquatica, Listeria cornellensis, be crucial in epidemiological investigations, source Listeria riparia, Listeria grandensis; Lang Halter tracking contamination from food processing plants ve ark. (25) Listeria weihenstephanensis; Weller ve and determining evolutionary relationships between ark. (47) ise Listeria booriae sp. nov. ve Listeria different strains. In this paper L. monocytogenes newyorkensis sp.nov. türlerini rapor etmişlerdir. Den strains and genetic and virulence differences among Bakker ve ark. (10) L. grayi’nin diğer Listeria türleri L. monocytogenes strains are reviewed. ile uzaktan akraba olduğunu ve farklı cins içine Keywords: Epidemiology, L. monocytogenes, alınmasını önermektedir. strains, virulence * Dr. Ziraat (Gıda) Müh., Sağlık Bakanlığı, Türkiye Halk Sağlığı Kurumu, Sıhhiye, Ankara. ** Prof. Dr., Ankara Üniversitesi, Veteriner Fakültesi, Gıda Hijyeni ve Teknolojisi AD, Dışkapı, Ankara. www.veteriner.org.tr/tr/dergi 97 Vet Hekim Der Derg 89(1): 97-107,2018 Çağrılı Makale / Invited Paper Orsi ve Wiedmann (33), Listeria cinsinin tüm L. newyorkensis, L. cornellensis, L. rocourtiae, L. üyelerinin fenotipik ve genotipik özellikleri ile weihenstephanensis, L. grandensis, L. riparia, L. ilgili son bilgileri inceledikleri çalışmalarında; bu booriae türlerini içermektedir. Orsi ve Wiedmann 17 türü, L. monocytogenes ile ilişkisini temel alarak (33), Listeria sensu lato içindeki 11 türü de üç farklı iki gruba ayırmıştır. (i) Listeria sensu strictu; L. monofiletik grupta açıklarken, ayrı cins olarak monocytogenes, L. marthii, L. innocua, L. ivanovii, tanınmasını önermektedirler (Şekil 1). L. welshimeri, L. seeligeri ve (ii) Listeria sensu lato; L. grayi, L. fleischmannii, L. floridensis, L. aquatica, T 100 Listeria monocytogenes ATCC 15313 95 Listeria marthii FSL$4 -1201T 100 Listeria innocua CLIP 11262T Listeria 100 Listeria welshimeri SLCC5334T 100 Listeria ivanovii PAM55 100 Listeria seeligerei SLCC 3954T T 100 Listeria grayi DSM 20601 Muraya Listeria grayi ATCC 25401T 95 100 Listeria fleichmanni subsp. fleiscmanni LU2006T 1 Listeria fleichmanni subsp. coloradonensis TTU M1 001T Mesolisteria T 100 Listeria floridonensis FSL S10-02091187 100 Listeria aquatica FSL S10-1088T 100 Listeria newyorkensis FSL A5-020 100 Listeria newyorkensis FSL M6-0635T 100 Listeria cornellensis TTU A1-0210 T 100 Listeria weihenstephanensis DSM 24699 T 100 Listeria grandensis TTU A1-012 Paenilisteria Listeria riparia FSL S10-1204T 100 Listeria booriae FSL A5-1281T T 100 Listeria rocourtiae CIP 109804 Listeria rocourtiae CIP 109804T 100 Brochothrix campestris ATOC 43754T Brochothrix thermosphacta ATCC 11509T Şekil 1: Listeria türlerinin filogenetik sınıflandırması ve önerilen yeni cins isimleri (33). Figure 1: Phylogenetic groups of Listeria species and proposed new genera names (33). Listeria sensu lato türlerinin hiçbiri patojen Listeria sensu lato grubundaki diğer türler flagellar değildir. Hem hemolitik testlerde hem de fosfoinosit proteinleri kodlayan hareket genlerini taşımaz. fosfolipaz C testlerinde pozitif değildirler. Ayrıca Flogenetik analizler, Listeria sensu strictu’nun genom analizleri Listeria sensu lato türlerinin; atası ve L. grayi ‘nin tüm flagellar biosentetik majör virülens genler prfA, plcA, hly, mpl, actA, genlerini, Basillus cereus kompleksinin atasından plcB genlerini içeren Listeria patojenite adası 1 horizantal gen transferi ile kazanıldığını göstermiştir. veya inlA, inlB içeren Listeria patojenite adası 2 Listeria sensu lato türleri, sensu strictu türleri ile bulundurmadığını göstermiştir. L. grayi dışında karşılaştırıldığında, internalin alanı ile ilgili genlerin 98 www.veteriner.org.tr/tr/dergi Vet Hekim Der Derg 89(1): 97-107,2018 Çağrılı Makale / Invited Paper yetersizliği görülmüştür. Bununla birlikte şaşırtıcı bazısının, değişen çevre şartlarına adaptasyon için biçimde, L. fleischmannii subsp. coloradonensis’in; gerekli elverişlilik faktörleri, diğerlerinin enfeksiyon Bacillus’dan MTX2 (putative mosquitocidal toxin) için gerekli virülens faktörler olduğu belirtilmektedir kodlayan genler bulundurduğu belirlenmiş ve bu (12). nedenle bazı Listeria sensu lato izolatlarının insektler L. monocytogenes, çevrede yaygın olarak gibi memeli olmayan konakçılarda hastalık nedeni bulunan ve gıdalarda kontrol edilmesi gereken bir olabileceği belirtilmiştir (33). patojendir. Bu nedenle Listeria monocytogenes Patojenik Listeria türlerinin her birinin, patojen insidensini azaltmak ve daha iyi kontrol önlemleri olmayan türlerle yakından ilişkili olduğu alabilmek için, mikroorganizma soylarının genetik ve virulens faktörlerinin belirlenmesi ve iyi anlaşılması belirtilmektedir. Patojen ve patojen olmayan Listeria gerekmektedir. Bu derlemede, etkene karşı alınacak türlerinin ve suşlarının farklılığının, yaklaşık 47 önlemlerin kolaylaştırılmasını sağlayabilecek milyon yıl önce, anahtar virülens genleri içeren faktörlerden olan L. monocytogenes soylarının patojenik atadan yaygınlaştığı; fakültatif patojenden genetik ve virulens farklılıklarının değerlendirilmesi saprofiteListeria türlerinin geçişinde gen kayıplarının amaçlanmıştır. kritik rolü olduğu belirtilmektedir. L. monocytogenes, L. monocytogenes’in soyları: Listeria türleri, L. innocua ve L. marthii ile; L. ivanovii, L. seeligeri Somatik (O faktör) ve Flagellar (H faktör) antijenlerine yakından ilişkilidir. L. seeligeri; Listeria patojenite göre tiplendirilmektedir (41). L. monocytogenes’in adası (LiPI) olarak da bilinen, temel Listeria virülens 1/2a, 1/2b, 1/2c, 3a, 3b, 3c, 4a, 4ab, 4b, 4c, 4d, gen bölgesinin benzerini içerse bile patojen olmadığı 4e ve 7 olmak üzere 13 serotipi vardır (13) Son belirtilmektedir (10). yıllarda, epidemiyolojik araştırmalarda ve kaynak Karşılaştırmalı genom çalışmalarında, L. izlemelerinde; yüksek ayrım gücü olan pulsed- monocytogenes’in 2853 geninin 270’inin (% field jel elektroforez (PFGE), multilocus sequence 10,5) L. innocua’da bulunmadığı belirlenmiştir. typing (MLST) gibi çeşitli genetik metodlar, alt L. monocytogenes CLIP80459 (serotip 4b) ile L. tiplendirme amacıyla kullanılmaktadır. Buna göre; monocytogenes EGDe (serotip 1/2a) suşlarının L. monocytogenes serotipleri genetik yapısına göre genleri karşılaştırıldığında ise CLIP80459 genlerinin önceleri 3 Soya (lineage) ayrılmış, Soy III de; IIIA, IIIB, IIIC olmak üzere 3 alt gruba ayrılmıştır. Daha yaklaşık % 8’nin L. monocytogenes EGDe sonra, Ward ve ark. (46), Soy IIIB’yi Soy 4 olarak genomunda olmadığı bildirilmiştir. L. monocytogenes ilk kez rapor etmişlerdir. Soy I; 1/2b, 3b, 4b, Soy izolatları arasındaki genetik farklılık, Listeria türleri II; 1/2a, 1/2c, 3c, Soy III ve IV; 4a, 4c ve atipik 4b arasındaki farklılığa yakındır. Bu farklılıklar yüzey serotiplerini içerir (Tablo 1) (32). proteinleri kodlayan genlerdeki farklılık ve şeker Liu ve ark. (28), Soy IIIA’nın tipik ramnoz pozitif metabolizmasında yer alan genlerdeki farklılıktan avirülent 4a serotipini ve virülent 4c serotipini, Soy kaynaklanmaktadır. L. monocytogenes’in 133 IIIC’nin; atipik ramnoz negatif virülent 4c serotipini, yüzey proteini kodlayan genin 30’unun (% 22,6), Soy IIIB’nin; atipik ramnoz negatif virülent olmayan 86 salgılanan proteininin 23’ünün (% 26,7) L. 4a ve virülent olmayan 4c serotipini içerdiğini innocua’da bulunmadığı belirlenmiştir. Bunların bildirmişlerdir. www.veteriner.org.tr/tr/dergi 99 Vet Hekim Der Derg 89(1): 97-107,2018 Çağrılı Makale / Invited Paper Tablo 1: L. monocytogenes soylarının özeti (32) Table 1: Summary of L. monocytogenes lineages (32) Soy İlk Tanımlama Serotip Genetik Özellikler Dağılım I Piffaretti ve ark. 1/2b,
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