唐崇惕,黃帥欽,彭午弦,盧明科,彭文峰,陳 東

湖北釘螺(Oncomelaniahupensis)被外睪吸蟲(Exorchisspp.)感染后很快產(chǎn)生大量血淋巴細(xì)胞和分泌物并能殺害再進(jìn)入其體內(nèi)的日本血吸蟲(Schistosomajaponicum)幼蟲[1,3,11]。外睪吸蟲和日本血吸蟲先后雙重感染釘螺間隔時(shí)間愈長(zhǎng)，血吸蟲被殺害的效果愈強(qiáng)，但釘螺血淋巴細(xì)胞的數(shù)目卻是逐漸減少[2,4]，釘螺體內(nèi)還有什么東西在起作用？近年許多學(xué)者在南美洲和非洲大力開展曼氏血吸蟲(Shistosomamansoni)與其中間宿主水生雙臍螺(Biomphalariaspp.)的相互關(guān)系問題的研究[5]。探討感染曼氏血吸蟲的雙臍螺體內(nèi)血漿中的蛋白質(zhì)組學(xué)(proteomics)，有關(guān)它們的多態(tài)性、多態(tài)粘蛋白(polymorphic mucins)[8-9],與血纖維源有關(guān)的蛋白質(zhì)(fibrinogen-related proteins) 等的特性、免疫應(yīng)答基因家族情況[13-14],有關(guān)軟體動(dòng)物防御細(xì)胞中激活酶(kinase)多方面作用情況[10]。有學(xué)者研究棘口吸蟲(Echinostomacaproni)和曼氏血吸蟲幼蟲在雙臍螺中的排泄和分泌的蛋白質(zhì)組(proteome)[7]，棘口吸蟲寄生的雙臍螺血漿及血淋巴細(xì)的不同蛋白質(zhì)的特性[6,12]。有關(guān)曼氏血吸蟲與雙臍螺相關(guān)的螺體血漿中物質(zhì)的研究報(bào)告無數(shù)，可見其中復(fù)雜程度。筆者從外睪吸蟲和日本血吸蟲雙重感染的釘螺體上觀察到有復(fù)雜的分泌物增多，它們?cè)诓煌g隔時(shí)間感染的釘螺體內(nèi)外表現(xiàn)情況有所不同，茲將觀察結(jié)果簡(jiǎn)單介紹于下。

1 材料與方法

用從湖南西洞庭湖鯰魚腸管收集的目平外睪吸蟲(ExorchismupingensisJiang,2011)的蟲卵拌以少量面粉飼食從西洞庭湖采集的湖北釘螺，外睪吸蟲感染后的釘螺分為5組，分別在感染后21 d、37 d、55 d、70 d、85 d，每粒釘螺與實(shí)驗(yàn)室日本血吸蟲陽(yáng)性小白鼠肝臟血吸蟲蟲卵孵化的毛蚴(40～90個(gè))接觸；各組釘螺均在感染血吸蟲毛蚴后4～82 d之間不同時(shí)間，用10%福爾馬林溶液固定。釘螺單獨(dú)感染外睪吸蟲后20 d，也用10%福爾馬林溶液固定。所有實(shí)驗(yàn)釘螺均經(jīng)石蠟連續(xù)切片及用蘇木精與洋紅染色制片。顯微鏡油鏡檢查各切片所有斷面，比較觀察各組無其它吸蟲天然感染的釘螺其體內(nèi)外分泌物的數(shù)量及它們的結(jié)構(gòu)情況，并數(shù)碼相機(jī)顯微照相儲(chǔ)存于電腦。

2 結(jié) 果

在不同間隔先后雙重感染目平外睪吸蟲和日本血吸蟲的釘螺體內(nèi)分泌物詳細(xì)情況的本實(shí)驗(yàn)觀察中，同樣見到間隔時(shí)間長(zhǎng)短與螺體殺害血吸蟲幼蟲的效力成正比，而與在螺體及異常血吸蟲幼蟲體內(nèi)的3種血淋巴細(xì)胞數(shù)量成反比的現(xiàn)象。但釘螺體內(nèi)外的分泌物密度沒有減少，有增多并且更加復(fù)雜化。情況分述如下。

2.1單獨(dú)感染外睪吸蟲的釘螺體表分泌物結(jié)構(gòu) 作為吸蟲類的中間宿主貝類，其體表都有一層分泌物。切片觀察單獨(dú)感染日本血吸蟲的釘螺，其體表的分泌物僅一層粘液性薄膜，而被外睪吸蟲感染的釘螺其體表分泌物增厚[11]。本實(shí)驗(yàn)觀察單獨(dú)感染外睪吸蟲后21 d的釘螺，其體表也有較厚的分泌物(圖1)，在油鏡下可見它們由排列不規(guī)則、大小和形狀各異、透明或不透明的晶體狀結(jié)構(gòu)所密布，其中及邊緣有些微粘液樣物質(zhì)(圖2～3)。

2.2釘螺體內(nèi)分泌物 在釘螺體中部、肉足和鰓的下方有一大團(tuán)副腺組織(圖6)，許多副腺細(xì)胞游離在螺體各組織中。在所有雙重感染外睪和血吸蟲的釘螺，副腺細(xì)胞亦可見于闊張的血管中[3]。在不同間隔雙重感染外睪和血吸蟲的釘螺，所有被擊毀的血吸蟲幼蟲殘?bào)w內(nèi)外都可見到這些細(xì)胞(圖7、9、22等)。

單獨(dú)感染外睪吸蟲的釘螺體內(nèi)血淋巴細(xì)胞和分泌物也比單獨(dú)感染日本血吸蟲的多，明顯的金黃色顆粒團(tuán)狀分泌物(圖4～5)出現(xiàn)在釘螺體內(nèi)一些區(qū)域。雙重感染外睪和血吸蟲的釘螺，雙重感染間隔時(shí)間加長(zhǎng)體內(nèi)分泌物有增無減，分泌物顆粒和血淋巴細(xì)胞經(jīng)常一起出現(xiàn)在被擊毀的異常血吸蟲幼蟲體內(nèi)外(圖8、10、13、17、19等)。有時(shí)它們會(huì)呈褐色團(tuán)狀大量地分布在螺體組織及異常血吸蟲體內(nèi)，以間隔55 d雙重感染后56 d(即外睪吸蟲感染后111 d，血吸蟲殘?bào)w為6 d)的照片(圖14)為例，大團(tuán)褐色物占據(jù)血吸蟲殘?bào)w大部，蟲體所在血腔外的螺體組織中密布許多小褐色團(tuán)。如此情況時(shí)?？梢?。

前此在外睪和血吸蟲雙重感染的釘螺體內(nèi)，已見到在金黃色分泌物顆粒每團(tuán)邊緣都有一個(gè)比小血淋巴細(xì)胞核更小的細(xì)胞核，其直徑只有1.9～2.6 μm[3]。此次用油鏡觀察，仍然是此情況，而且在核的外圍都可見到有一圈白色外圍，它應(yīng)該是其細(xì)胞質(zhì)部位。說明它們確有細(xì)胞的結(jié)構(gòu)，雙重感染時(shí)間愈長(zhǎng)此極小細(xì)胞愈明顯(圖8、11～12、16、18～20)。

2.3釘螺體內(nèi)分泌物顆粒在異常血吸蟲幼蟲體內(nèi)的異樣產(chǎn)物 在外睪和血吸蟲雙重感染的釘螺體內(nèi)，尚有奇怪現(xiàn)象：在很早期被擊毀的異常血吸蟲幼蟲體內(nèi)會(huì)出現(xiàn)一個(gè)大紅色球團(tuán)，它整個(gè)表面布滿顆粒小點(diǎn)，球團(tuán)邊緣圍繞甚多與上述分泌物顆粒上極小細(xì)胞同樣大小的胞核和細(xì)胞樣的結(jié)構(gòu)(圖8～10、17)。雙重感染間隔時(shí)間愈長(zhǎng)的釘螺，異常血吸蟲幼蟲體內(nèi)此紅球團(tuán)數(shù)常會(huì)增到2～4個(gè),而且歷久不退，如間隔70 d后的6 d和25 d異常血吸蟲幼蟲的樣品(圖15、21)。

2.4外睪吸蟲和日本血吸蟲雙重感染的釘螺體表分泌物結(jié)構(gòu) 外睪吸蟲和日本血吸蟲雙重感染的釘螺，它們體表分泌物均有增無減，其厚度可達(dá)單獨(dú)感染外睪吸蟲釘螺體表分泌物厚度數(shù)倍(圖23)。兩吸蟲感染間隔時(shí)間增長(zhǎng)，其體表分泌物不僅增多，而且其內(nèi)容更加復(fù)雜化。其中晶體狀物體逐漸呈有規(guī)則地成條成片地排列(圖26～27)，在許多晶體樣物質(zhì)之間除了有更多粘液樣物體之外還有許多原本是在釘螺體內(nèi)的物質(zhì)。兩吸蟲間隔85 d雙重感染，在血吸蟲感染后5 d～25 d的釘螺，它們體中都只能查到已完全解體的血細(xì)蟲幼蟲殘骸。在血吸蟲感染后5 d的釘螺體表分泌物(圖23)中在許多成片條狀晶體物質(zhì)之間除有許多粘液狀物質(zhì)之外還夾雜著許多原來在釘螺體內(nèi)的血淋巴細(xì)胞和含極小細(xì)胞的分泌物顆粒(圖24～28)。

圖版箭矢數(shù)字說明(Arrownumbersillustrate):

1=大血淋巴細(xì)胞(bighemo-lymphocyte)；2=中血淋巴細(xì)胞(mediatebighemo-lymphocyte)；

3=小血淋巴細(xì)胞(smallhemo-lymphocyte)；4=螺體外分泌物(snailbodyoutersecretion)；

5=螺體內(nèi)分泌物及小核(snailinnersecretionandverysmallnucleus)；

6=螺副腺細(xì)胞(snailaccessoryglandcell)。

圖說明(Em=Exorchismupingensis;Oh=Oncomelaniahupensis;Sj=Schistosomajaponicum)

圖1～6被Em感染的Oh其分泌物及副腺細(xì)胞

Figs.1-6SecretionsandaccessoryglandcellsofOhsnailsinfectedbyEmtrematodes

圖7間隔21d雙重感染Em和Sj的Oh示其體內(nèi)分泌物和血淋巴細(xì)胞，及其對(duì)Sj幼蟲的反應(yīng)

Fig.7OhsnailduallyinfectedbyEmandSjfor21dinterval,showingthesnail’ssecretions,lymphocytesandtheirreactionstoSjlarvae

圖8～11.間隔37d雙重感染Em和Sj的Oh示其體內(nèi)分泌物和血淋巴細(xì)胞，及其對(duì)Sj幼蟲的反應(yīng)

Figs.8-11OhsnailsduallyinfectedbyEmandSjfor37dinterval,showingthesnail’ssecretions,lymphocytesandtheirreactionstoSjlarvae

圖12～14間隔55d雙重感染Em和Sj的Oh示其體內(nèi)分泌物和血淋巴細(xì)胞，及其對(duì)Sj幼蟲的反應(yīng)

Figs.12-14OhsnailsduallyinfectedbyEmandSjfor55dinterval,showingthesnail’ssecretions,lymphocytesandtheirreactionstoSjlarvae

圖15～22間隔70d雙重感染Em和Sj的Oh示其體內(nèi)分泌物和血淋巴細(xì)胞，及其對(duì)Sj幼蟲的反應(yīng)

Figs.15-22OhsnailsduallyinfectedbyEmandSjfor70dinterval,showingthesnail’ssecretions,lymphocytesandtheirreactionstoSjlarvae

圖23～28.間隔85d雙重感染Em和Sj的Oh示其體內(nèi)分泌物和血淋巴細(xì)胞，及其對(duì)Sj幼蟲的反應(yīng)

Figs.23-28OhsnailsduallyinfectedbyEmandSjfor85dinterval,showingthesnail’ssecretions,lymphocytesandtheirreactionstoSjlarvae

圖1Oh體外分泌物 (Scale bar=0.15 mm)

Fig.1 Outer secretions ofOhsnail

圖2Oh體外分泌物 (Scale bar=0.03 mm)

Fig.2 Outer secretions ofOhsnail

圖3Oh體外分泌物(Scale bar=0.023 mm)

Fig.3 Outer secretions ofOhsnail

圖4Oh體內(nèi)分泌物(Scale bar=0.034 mm)

Fig.4 Inner secretions ofOhsnail

圖5Oh體內(nèi)分泌物(Scale bar=0.030 mm)

Fig.5 Inner secretions ofOhsnail

圖6Oh體內(nèi)副腺細(xì)胞(Scale bar=0.022 mm)

Fig.6 Accessory gland cells ofOhsnail

圖7Em感染后37 dOh體中3種血淋巴細(xì)胞在16 d異常Sj幼蟲體內(nèi)外(Scale bar=0.034 mm)

Fig.7 Three species hemo-lymphocytes in 16-d-old abnormalSjlarva and tissue of snail post-infected byEmfor 37 d

圖8 含極小細(xì)胞核的分泌顆粒散布在感染Em后42 d的Oh體組織中，具分泌物顆粒的球形紅團(tuán)在5d異常Sj幼蟲體內(nèi)(Scale bar=0.022 mm)

Fig.8 The secretion granules with very small cell nucleus spreading in the tissue ofOhpost-infected byEmfor 42 d and a spherical red mass with secretions granules in 5-d-old abnormalSjlarva

圖9Em感染后42 d的Oh示其體中5d異常Sj幼蟲含副腺細(xì)胞及具分泌物顆粒的紅團(tuán)(Scale bar=0.027 mm)

Fig.9 Body ofOhpost-infected byEmfor 42 d showing the 5-d-old abnormalSjlarva containing the accessory gland cells and a spherical red mass with secretions granules

圖10Em感染后42 d的Oh示其體中血淋巴細(xì)胞及具紅團(tuán)的5d異常Sj幼蟲 (Scale bar=0.030 mm)

Fig.10 Body ofOhpost-infected byEmfor 42 d showing the hemo-lymphocytes and 5-d-old abnormalSjlarva with hemo-cytes,red mass and secretion granules

圖11 帶有微小細(xì)胞核的分泌物顆粒團(tuán)散布在Em感染后42 d的Oh組織中 (Scale bar=0.028 mm)

Fig.11 Secretion granules with small cell nucleus spreading in the tissue ofOhpost-infected byEmfor 42 d

圖12Em感染后66 d 的Oh和11 d 異常Sj幼蟲體中的具微小細(xì)胞核的分泌物顆粒(Scale bar=0.030 mm)

Fig.12 Secretion granules with small cell nucleus inOhpost-infected byEmfor 66 d and the body of 11-d-old abnormalSjlarva

圖13Em感染后79 d的Oh示24 d 異常Sj幼蟲殘骸內(nèi)的血淋巴細(xì)胞和具微小細(xì)胞核的分泌物顆粒 (Scale bar=0.025 mm)

Fig.13Ohpost-infected byEmfor 79 d showing the hemo-lymphocytes and secretion granules with small cell nucleus in the 24-d-old abnormalSjlarva wreckage

圖14 大量分泌物團(tuán)在Em感染后111 d的Oh及56 d異常Sj幼蟲殘骸內(nèi) (Scale bar=0.033 mm)

Fig.14Much secretions masses inOhpost-infected byEmfor 111 d and the 56-d-old abnormalSjlarva wreckage

圖15Em感染后76 d的Oh示6 d異常Sj幼蟲殘骸內(nèi)的具分泌物顆粒2個(gè)紅團(tuán)(Scale bar=0.016 mm)

Fig.15Ohpost-infected byEmfor 76 d showing two red masses with secretion granules and small cell nucleuses in the 6-d-old abnormalSjlarva wreckage

圖16 間隔70 d雙重感染Em和Sj后6 d的Oh體中散布許多具微小細(xì)胞核的分泌顆粒(Scale bar=0.028 mm)

Fig.16 Many secretion granules with small cell nucleus in theOhat the 6 d after dual infections ofEmandSjwith 70 d intervals

圖17Em感染后84 d的Oh示14 d異常Sj幼蟲殘骸內(nèi)的紅團(tuán)和微小細(xì)胞核(Scale bar=0.027 mm)

Fig.17Ohpost-infected byEmfor 84 d showing a red mass with small cell nucleuses in the 14-d-old abnormalSjlarva wreckage

圖18 間隔70 d雙重感染Em和Sj后25 d的Oh體中的許多具微小細(xì)胞核的分泌顆粒 (Scale bar=0.027 mm)

Fig.18 Many secretion granules with small cell nucleus in theOhat the 25 d after dual infections ofEmandSjwith 70 d intervals

圖19Em感染后84 d的Oh示25 d異常Sj幼蟲殘骸內(nèi)外的具微小細(xì)胞核分泌物顆粒 (Scale bar=0.028 mm)

Fig.19Ohpost-infected byEmfor 84 d showing the secretion granules with small cell nucleus inside and outside of 25d old abnormalSjlarva wreckage

圖20 間隔70 d雙重感染Em和Sj后25 d的Oh體內(nèi)許多具微小細(xì)胞核的分泌顆粒 (Scale bar=0.025 mm)

Fig.20Many secretion granules with small cell nucleus in theOhat the 25 d after dual infections ofEmandSjwith 70 d intervals

圖21Em感染后95 d的Oh示2個(gè)具分泌物顆粒的紅團(tuán)在25 d 異常Sj幼蟲殘骸內(nèi) (Scale bar=0.027 mm)

Fig.21Ohpost-infected byEmfor 95 d showing two red masses with secretion granules in the 25-d-old abnormalSjlarva wreckage

圖22Em感染后95 d的Oh示副腺細(xì)胞在25d異常Sj幼蟲殘骸內(nèi) (Scale bar=0.030 mm)

Fig.22Ohpost-infected byEmfor 95 d showing the accessory gland cells in the 25-d-old abnormalSjlarva wreckage

圖23 間隔85 d雙重感染Em和Sj后5 d的Oh示體外分泌物(Scale bar=0.15 mm)

Fig.23Ohat the 5 d after dual infections ofEmandSjwith 85 d intervals showing its thick body secretion

圖24 間隔85 d雙重感染Em和Sj后5 d的Oh示體外分泌物中的體內(nèi)血淋巴細(xì)胞(Scale bar=0.027 mm)

Fig.24Ohat the 5 d after dual infections ofEmandSjwith 85 d intervals showing hemo-lymphocytes in the body outside secretion

圖25 間隔85d雙重感染Em和Sj后5 d的Oh示體外分泌物中的體內(nèi)小血淋巴細(xì)胞和分泌物顆粒微小細(xì)胞核(Scale bar=0.027 mm)

Fig.25Ohat the 5 d after dual infections ofEmandSjwith 85 d intervals showing small hemo-lymphocytes and secretion granules with small cell nucleus in the body outside secretion

圖26 間隔85 d雙重感染Em和Sj后5 d的Oh示體外分泌物中的結(jié)晶體和體內(nèi)分泌物顆粒微小細(xì)胞核(Scale bar=0.019 mm)

Fig.26Ohat the 5 d after dual infections ofEmandSjwith 85 d intervals showing the opaque crystal structure of body secretion and inner secretion granules with small cell nucleus in the body outside secretion

圖27 間隔85 d雙重感染Em和Sj后5 d的Oh示體外分泌物中的結(jié)晶體和體內(nèi)分泌物顆粒微小細(xì)胞核(Scale bar=0.019 mm)

Fig.27Ohat the 5 d after dual infections ofEmandSjwith 85 d intervals showing the opaque crystal structure of body secretion and inner secretion granules with small cell nucleus in the body outside secretion

圖28 間隔85 d雙重感染Em和Sj后5 d的Oh的體外分泌物中的結(jié)晶體和體內(nèi)分泌物顆粒微小細(xì)胞核(Scale bar=0.019 mm)

Fig.28Ohat the 5 d after dual infections ofEmandSjwith 85 d intervals showing the opaque crystal structure of body secretion and inner secretion granules with small cell nucleus in the body outside secretion

3 討 論

實(shí)驗(yàn)證明外睪吸蟲和日本血吸蟲雙重感染釘螺，其間隔時(shí)間愈長(zhǎng)螺體殺害血吸蟲幼蟲的效力愈強(qiáng)，釘螺感染外睪吸蟲的開始時(shí)會(huì)增生大量血淋巴細(xì)胞，但在感染后一個(gè)月左右螺體增生血淋巴細(xì)胞性能就開始逐漸衰退，時(shí)間稍久后其數(shù)量逐漸顯著減少[2,4]；外睪吸蟲幼蟲期在釘螺體內(nèi)無性生殖期很長(zhǎng)，感染后105 d還處于原始胚細(xì)胞大量增殖期，6～7個(gè)月才成熟[1]。本實(shí)驗(yàn)外睪吸蟲陽(yáng)性釘螺同樣表現(xiàn)在其感染后37～85 d能逐漸地更有力地攻擊后侵入的血吸蟲幼蟲，使其致命解體，是否與螺體內(nèi)外睪吸蟲胚細(xì)胞正在大量增生有關(guān)？螺體內(nèi)分泌物逐漸大量增加是否也與其有關(guān)？但是為何螺體的血淋巴細(xì)胞卻在逐漸減少？其中奧秘機(jī)理需要繼續(xù)探究。

釘螺體內(nèi)外的分泌物從那里產(chǎn)生？在外睪吸蟲和日本血吸蟲雙重感染的釘螺其體外分泌物中呈晶體結(jié)構(gòu)是何物質(zhì)？體內(nèi)金黃色分泌物顆粒有增無減，每個(gè)顆粒團(tuán)都更明顯出有細(xì)胞核和細(xì)胞質(zhì)的極小細(xì)胞，這細(xì)胞和金黃色顆粒團(tuán)是何關(guān)系？這些分泌物與血吸蟲幼蟲被擊毀有何關(guān)系？其中機(jī)理亦需繼續(xù)探究。

外睪吸蟲和日本血吸蟲雙重感染的釘螺，其體中被殺害的血吸蟲幼蟲體內(nèi)經(jīng)常有球狀紅團(tuán)，兩吸蟲雙重感染間隔時(shí)間愈長(zhǎng)這紅團(tuán)數(shù)會(huì)從1個(gè)增加到4個(gè)。這紅團(tuán)的產(chǎn)生與其外表分泌物顆粒及極小細(xì)胞核相仿的物體有何關(guān)系？與血吸蟲幼蟲的被殺有無關(guān)系？其中機(jī)理亦需繼續(xù)探究。

應(yīng)用無害的外睪吸蟲作材料處理釘螺，所有實(shí)驗(yàn)都證明可以百分之百殺死再侵入的日本血吸蟲幼蟲，有關(guān)此生物控制的機(jī)制都需要從多方面，包括免疫學(xué)、蛋白質(zhì)組學(xué)和基因組學(xué)等等，進(jìn)行深入研究，有利于更好地應(yīng)用。當(dāng)時(shí)研究生郭躍和王逸難同學(xué)協(xié)助感染工作，在此致謝。

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