李十盛,高 會,趙富強,陳以芹,張克玉,黃加勁,那廣水,3*

水產(chǎn)養(yǎng)殖環(huán)境中抗生素抗性基因的研究進展

李十盛1,2,高 會2**,趙富強2,4,陳以芹2,張克玉1,2,黃加勁1,2,那廣水1,2,3*

(1.上海海洋大學海洋生態(tài)與環(huán)境學院,上海市 201306；2.國家海洋環(huán)境監(jiān)測中心,遼寧 大連 116023；3.海南熱帶海洋學院生態(tài)環(huán)境學院,海南 三亞 572022；4.大連海洋大學海洋科技與環(huán)境學院,遼寧 大連 116023)

為摸清我國水產(chǎn)養(yǎng)殖環(huán)境抗生素抗性基因(Antibiotic resistance genes, ARGs)污染狀況,統(tǒng)計了當前水產(chǎn)養(yǎng)殖環(huán)境水、沉積物和養(yǎng)殖對象中七大類抗性基因的賦存狀況,進一步分析了影響其賦存的因素,對未來我國水產(chǎn)養(yǎng)殖ARGs污染研究進行了展望.水產(chǎn)養(yǎng)殖環(huán)境中ARGs主要以iDNA形式存在;磺胺類和四環(huán)素類抗性基因的豐度和檢出率較高,被認為是水產(chǎn)養(yǎng)殖環(huán)境主要的ARGs,已受到當前研究的廣泛關(guān)注;而喹諾酮類、氯霉素類、β-內(nèi)酰胺類、大環(huán)內(nèi)酯類和氨基糖苷類抗性基因報道的豐度和檢出率相對較低.養(yǎng)殖過程中,養(yǎng)殖用藥、飼料及養(yǎng)殖模式是影響水產(chǎn)養(yǎng)殖環(huán)境ARGs賦存的主要因素,但環(huán)境理化因子調(diào)節(jié)養(yǎng)殖環(huán)境ARGs的作用也不容忽視.

抗生素抗性基因(ARGs)；水產(chǎn)養(yǎng)殖環(huán)境；賦存狀況；影響因素

我國水產(chǎn)養(yǎng)殖養(yǎng)殖總面積高達719萬hm2[1],水產(chǎn)養(yǎng)殖產(chǎn)量占全球的61.2%[2].我國水產(chǎn)養(yǎng)殖用藥的種類繁多,常用的藥品有抗菌類藥物、水體消毒劑、抗寄生蟲藥物、抗真菌藥物等九類[3].其中我國批準應用于水產(chǎn)養(yǎng)殖的抗菌類藥物包括多甲砜霉素、四環(huán)素、恩諾沙星、氟苯尼考、磺胺嘧啶和甲氧芐啶[4]等數(shù)十種.但隨著水產(chǎn)養(yǎng)殖產(chǎn)業(yè)的飛速發(fā)展,為了提高水產(chǎn)養(yǎng)殖對象產(chǎn)量,降低養(yǎng)殖成本,抗生素作為養(yǎng)殖藥物或混合飼料大量投入使用,這和健康養(yǎng)殖相互矛盾[5].水產(chǎn)養(yǎng)殖環(huán)境抗生素被認為是自然水體抗生素主要來源之一[6-7],如長江流域和珠江下游抗生素濃度在N.D.～537ng/L之間[8-9];海河流域抗生素平均濃度為821ng/L[10];南黃海和東海沿岸抗生素濃度在N.D.～497.6ng/L之間[6,11],北部灣河口和沿海抗生素總濃度為1.81～118ng/L[7];黃海、渤?？股貪舛葍HN.D.～16.6ng/L[12].而黃海周邊典型水產(chǎn)養(yǎng)殖場和中國東南部海水養(yǎng)殖場中抗生素濃度在N.D.～ 995.02ng/L[13-14];北部灣海水養(yǎng)殖區(qū)抗生素總濃度達43.2～885ng/L,遠高于其周圍環(huán)境水體[7].抗生素的頻繁使用不僅降低了抗生素的藥效,還促進水產(chǎn)養(yǎng)殖環(huán)境和養(yǎng)殖生物體內(nèi)抗性細菌及抗生素抗性基因(Antibiotic resistance genes, ARGs)的發(fā)生,加速了ARGs的傳播.導致水產(chǎn)養(yǎng)殖環(huán)境成為抗生素和ARGs的潛在儲藏庫[14-15].

ARGs是可使細菌能耐受更高濃度抗生素,或者其缺失會導致細菌對抗生素的易感性增加的基因,其可原本便存在于環(huán)境細菌的基因組中,也可細菌經(jīng)過基因突變而獲得[16],通過垂直基因轉(zhuǎn)移[17]和水平基因轉(zhuǎn)移[18-19]傳播.但養(yǎng)殖環(huán)境中持續(xù)的選擇壓力(抗生素等),加速了ARGs出現(xiàn)、入侵和侵占這一過程[20].如果抗生素選擇壓力足夠大,很低概率的抗性也可被細菌選擇獲得和轉(zhuǎn)移[21].Davies等[22]表示ARGs污染及其在整個生物圈微生物種群中的分布是人類使用抗生素的持續(xù)選擇的結(jié)果,這不是自然過程,而是疊加在自然上的人為情況.

水產(chǎn)養(yǎng)殖環(huán)境中,ARGs隨著菌群遷移傳播,可直接進入養(yǎng)殖生物體內(nèi),最終通過食物鏈和人類腸道微生物群進行水平基因轉(zhuǎn)移[15,23-24].Yang等[25]發(fā)現(xiàn)海水養(yǎng)殖沉積物中質(zhì)粒和轉(zhuǎn)座子攜帶的ARGs與人類致病菌中的ARGs序列高度相似.即使攜帶ARGs的細菌被殺滅,它仍可以釋放到環(huán)境中,并隨著養(yǎng)殖水體擴散或定期交換到自然水體當中[26],嚴重影響生態(tài)健康.Hedberg等[27]研究也表明水產(chǎn)養(yǎng)殖尾水是沿海水域ARGs的重要來源.由于ARGs在環(huán)境和生物體內(nèi)的傳播長久而持續(xù),造成生態(tài)和食品安全的不確定性風險,引發(fā)社會越來越多的關(guān)注.世界衛(wèi)生組織(WHO)將抗生素耐藥性的傳播列為21世紀公共衛(wèi)生面臨的三大最嚴重威脅之一[28].ARGs作為一類新型環(huán)境污染物,其在水產(chǎn)養(yǎng)殖環(huán)境介質(zhì)中的傳播擴散可能比抗生素本身的危害更大[29].

與醫(yī)療[30-31]、污水處理[32-33]、禽畜糞便[34-35]等環(huán)境相比,水產(chǎn)養(yǎng)殖環(huán)境由于養(yǎng)殖模式、養(yǎng)殖品種、養(yǎng)殖階段等不同,甚至城市污水處理廠、畜禽養(yǎng)殖場未被完全清除的抗生素及ARGs排放到河流[36-38],隨著水體交換最終進入水產(chǎn)養(yǎng)殖環(huán)境,導致水產(chǎn)養(yǎng)殖中ARGs的研究及控制更難以開展.水產(chǎn)養(yǎng)殖環(huán)境中的ARGs作為影響人類健康、生態(tài)系統(tǒng)、養(yǎng)殖生產(chǎn)力重要節(jié)點[39](圖1),而其賦存狀況及影響因素研究相對較少.因此,本文以水產(chǎn)養(yǎng)殖環(huán)境為研究區(qū)域,結(jié)合近年相關(guān)文獻(以PCR檢測方法為基礎(chǔ)),分析了水產(chǎn)養(yǎng)殖環(huán)境中喹諾酮類、β-內(nèi)酰胺類、大環(huán)內(nèi)酯類、氯霉素類、四環(huán)素類、磺胺類(含甲氧芐氨嘧啶類)以及氨基糖苷類ARGs賦存狀況,探究了影響水產(chǎn)養(yǎng)殖環(huán)境ARGs賦存和傳播的影響因素,為未來水產(chǎn)養(yǎng)殖環(huán)境ARGs的深入研究和控制提供科學依據(jù).

圖1 抗生素、抗性菌和ARGs在水產(chǎn)養(yǎng)殖環(huán)境中作用的因果關(guān)系

1 水產(chǎn)養(yǎng)殖環(huán)境中抗生素抗性基因賦存狀況

1.1 水產(chǎn)養(yǎng)殖環(huán)境中抗生素抗性基因存在形態(tài)

目前環(huán)境中的ARGs主要以兩種形式存在: iARGs和eARGs.iARGs是胞內(nèi)DNA(iDNA)所攜帶的ARGs,主要存在抗生素抗性細菌體內(nèi),可以通過復制傳給下一代,也可以通過接合和轉(zhuǎn)導傳遞給其它物種[8,40].eARGs是游離態(tài)的胞外DNA(eDNA)所攜帶ARGs,可由死細菌裂解或活細菌分泌釋放,裸露于環(huán)境介質(zhì)中,并具有很長的存在周期,可通過轉(zhuǎn)化方式進入到水環(huán)境的細菌中,使細菌獲得抗生素抗性[8,40].與iARGs相比,eARGs因為其特有的性質(zhì)更容易在環(huán)境中傳播,對人體健康和生態(tài)系統(tǒng)產(chǎn)生嚴重的威脅,研究表明[41]豐富的eDNA導致了水生環(huán)境中ARGs高轉(zhuǎn)移頻率.可見,胞內(nèi)外ARGs的傳播擴散形式大不相同.

研究發(fā)現(xiàn)在渤海海水,來自醫(yī)院、制藥廠、污水處理廠的污泥,以及渤海、海河、城市湖泊的沉積物中,eARGs的豐度皆高于iARGs,沉積物中的eDNA被認為可被粘土礦物、沙子和腐殖質(zhì)吸收,保護它不受部分核酸酶失活和物理分離的攻擊[19,42-43].與上述區(qū)域不同,Yuan等[41]在汕頭的牛蛙和魚,蝦和蟹混養(yǎng)的養(yǎng)殖場沉積物中發(fā)現(xiàn),ARGs主要以iARGs形式存在,iDNA中可檢測ARGs的平均檢測頻率(60.3%)高于eDNA(40.6%).同時,在牲畜糞便中,eDNA占污泥中總DNA的不到1.5%[35],與汕頭養(yǎng)殖場的ARGs的胞內(nèi)胞外分布相似.這可能是eDNA主要來源是死細胞的裂解釋放[42],而牲畜和水產(chǎn)養(yǎng)殖場的營養(yǎng)豐富,能夠很好地支持活細胞的生長和存活.因此, iDNA可能是水產(chǎn)養(yǎng)殖環(huán)境中ARGs的主要存在形式.

1.2 水產(chǎn)養(yǎng)殖環(huán)境水體中抗生素抗性基因賦存狀況

ARGs在水產(chǎn)養(yǎng)殖環(huán)境水體中普遍存在,ARGs相對豐度水平在1.28×10-8～1.87×10-1copies/16s rRNA之間(見表1).目前水產(chǎn)養(yǎng)殖環(huán)境水體中ARGs研究集中在磺胺類(包括甲氧芐氨嘧啶)、四環(huán)素類、喹諾酮類、氯霉素類、β-內(nèi)酰胺類、大環(huán)內(nèi)酯類六大類.其中磺胺類ARGs(含甲氧芐氨嘧類)有;四環(huán)素類ARGs有;喹諾酮類ARGs有;氯霉素類ARGs有;β-內(nèi)酰胺類ARGs有;大環(huán)內(nèi)酯類ARGs有(見圖2).

在多數(shù)報道中,磺胺類豐度占養(yǎng)殖水總ARGs相對最高,被認為是中國海水養(yǎng)殖場中最常檢測到的抗性基因[14].其次是四環(huán)素類ARGs,其亞型最為豐富,和在韓國海水養(yǎng)殖場可占總ARG的74.8%～98.0%[44].但是不同養(yǎng)殖區(qū)域,養(yǎng)殖對象不同造成的ARGs賦存狀況存在很大差異.如喹諾酮類在珠江三角洲河口白腿蝦養(yǎng)殖區(qū)為主要檢出ARG[45];在杭州灣河口蝦養(yǎng)殖區(qū)相對豐度能達到9.97×10-3copies/16s rRNA[46],這可能是喹諾酮類抗生素在蝦養(yǎng)殖中大量使用,且喹諾酮類ARGs在沿海水域研究[47]中是主要的抗性基因.而氯霉素類ARGs,在數(shù)地[44-46,48]均有報道;水東灣蝦場蝦塘水中有極高豐度[48].β-內(nèi)酰胺類ARGs的研究報道相對較少卻不容忽視,韓國沿海養(yǎng)殖場相對豐度能達到1.04×10-3copies/16s rRNA[44,49].

表1 水產(chǎn)養(yǎng)殖環(huán)境水體中ARGs賦存

續(xù)表1

注:a為相對豐度(copies/16S rRNA);b為絕對豐度(copies/mL;copies/g);-為數(shù)據(jù)未明確報道;( )內(nèi)為檢出率;加粗為報道主要ARG.表2、3同上.

1.3 水產(chǎn)養(yǎng)殖環(huán)境沉積物中抗生素抗性基因賦存狀況

據(jù)統(tǒng)計水產(chǎn)養(yǎng)殖環(huán)境沉積物中ARGs相對豐度在1.25×10-6～1.08×10-1copies/16s rRNA之間(見表2).其中磺胺類ARGs(含甲氧芐氨嘧類)有、、、/、;四環(huán)素類ARGs有、、、、、、、、、、、、、、、-;喹諾酮類ARGs有、、、;氯霉素類ARGs有、、、、_、_、;β-內(nèi)酰胺類ARGs有、-、-、-、、;大環(huán)內(nèi)酯類ARGs有、、;氨基糖苷類ARGs有、、(見圖2).、豐度依然較高,被認為是珠江口水產(chǎn)養(yǎng)殖[45]沉積物中ARGs的潛在指標;被認為是珠江三角洲養(yǎng)殖場沉積物中四環(huán)素ARGs豐度的潛在指標[49].

表2 水產(chǎn)養(yǎng)殖環(huán)境沉積物中ARGs賦存

與水不同,沉積物是ARGs的一個重要的“匯”,且沉積物中ARGs存在持久性.越南集約化蝦養(yǎng)殖區(qū)[53]、海南東寨港[50]等地沉積物中的ARGs的賦存量均大于水.Yuan等[46]發(fā)現(xiàn)隨著時間的推移,ARGs的擴散導致其可在沉積物中積累,沉積物很可能增強抗生素抗性的傳播.但有研究[2]認為ARGs在沉積物群落中的“持久性”,是水生生物糞便中ARGs的不斷涌入所導致的.此外,不同的ARGs在水和沉積物中存在不同的賦存規(guī)律.統(tǒng)計發(fā)現(xiàn)如等ARGs僅在沉積物有報道,()()--等卻僅在水中有報道(圖2).Su等[48]研究也證實了在水產(chǎn)養(yǎng)殖環(huán)境中,ARGs的賦存在不同介質(zhì)中有明顯差異.不過,PCR檢測的目標ARGs的局限性也可造成水和沉積物中ARGs統(tǒng)計結(jié)果產(chǎn)生差異,在煙臺海水養(yǎng)殖場[54]、瀛湖養(yǎng)殖區(qū)[55]運用宏基因組方法及高通量PCR方法檢出的ARGs則更加全面.

圖2 近年水產(chǎn)養(yǎng)殖環(huán)境中ARGs檢出報道頻次熱

1.4 水產(chǎn)養(yǎng)殖對象中抗生素抗性基因賦存狀況

水產(chǎn)養(yǎng)殖中抗生素主要作用于養(yǎng)殖生物,產(chǎn)生耐藥性不可避免.當前對水產(chǎn)養(yǎng)殖生物的ARGs的研究主要針對于其本身以及其腸道菌群.而鑒于運用PCR檢測方法研究水產(chǎn)養(yǎng)殖生物ARGs的報道相對較少,本文結(jié)合了運用宏基因組技術(shù)的相關(guān)研究.在水產(chǎn)養(yǎng)殖生物中,喹諾酮類、β-內(nèi)酰胺類、大環(huán)內(nèi)酯類、氯霉素類、四環(huán)素類、磺胺類、氨基糖苷類七大類ARGs皆有報道,其中被報道頻率較高(見圖2、表3).特別的,桿菌肽類、多重耐藥性和其他類ARGs(如等)豐度非常高[24,60].桿菌肽鋅作為禁用的飼料添加劑,其ARGs似仍以很高的量存在,但未發(fā)現(xiàn)有運用PCR技術(shù)針對性對其檢測的報道.而多重耐藥性和其他類ARGs的功能和特性復雜,可促使“超級細菌”形成,對人類健康及生態(tài)環(huán)境威脅更大.但目前對多重耐藥性ARGs的認識還相對較淺,其種類繁多,相關(guān)數(shù)據(jù)庫有待豐富,需更多研究作為基礎(chǔ).

表3 水產(chǎn)養(yǎng)殖生物體中ARGs賦存

續(xù)表3

值得注意的是,抗生素在水生生物體內(nèi)經(jīng)食物鏈富集已經(jīng)得到重視,但水產(chǎn)養(yǎng)殖環(huán)境和養(yǎng)殖生物體內(nèi)賦存的ARGs之間的聯(lián)系的研究還處在起步階段.早在2003年,Furushita等[61]就發(fā)現(xiàn)在魚類細菌中檢測到的四環(huán)素耐藥基因與人類臨床分離株中的基因高度相似.當前認為水平基因轉(zhuǎn)移是ARGs水體、沉積物和水產(chǎn)品之間轉(zhuǎn)移的關(guān)鍵途徑[8,41,52],且可移動遺傳元件(Mobile Genetic Element,MGEs)對ARGs轉(zhuǎn)移起著重要作用[29,62],其本身可攜帶ARGs,也與ARGs有共發(fā)生現(xiàn)象,往往代表著ARGs的高遷移率.Su等[48]通過Source Tracer分析發(fā)現(xiàn)早期和中期的蝦腸道約70%的菌群起源于沉積物,且蝦腸道和沉積物賦存了相同的ARGs().由此,賦存在水產(chǎn)養(yǎng)殖水體和沉積物中的ARGs很可能最終富集到水產(chǎn)養(yǎng)殖對象的體內(nèi).

2 水產(chǎn)養(yǎng)殖環(huán)境中抗生素抗性基因的影響因素

2.1 養(yǎng)殖用藥及飼料對抗生素抗性基因的影響

養(yǎng)殖用藥及飼料的使用對ARGs發(fā)生及傳播的影響一直被密切關(guān)注.對于養(yǎng)殖用藥,ARGs在養(yǎng)殖水域大量存在和藥物在治療養(yǎng)殖生物疾病中大量使用是分不開的.ARGs與相應的同源抗生素顯著相關(guān)[66].從機制上,長期處于養(yǎng)殖用藥的環(huán)境中,細菌的SOS反應(由細菌DNA損傷引發(fā)的修補反應)會被激活,其可以通過產(chǎn)生氧離子而增加突變幾率,促使水平基因位移,活化整合酶,促使整合子重組,從而顯著增強ARGs的水平轉(zhuǎn)移[56-67].控制水產(chǎn)養(yǎng)殖中抗生素類用藥是減緩ARGs污染的重要手段.而對于飼料,市場上用于養(yǎng)殖水產(chǎn)品的飼料種類繁多,?；旌峡股厥褂?甲殼類水產(chǎn)品(日本沼蝦)長期暴露在飼料中抗生素(合法使用量)下,也會嚴重影響蝦腸道健康和ARGs豐度[68].其中,魚粉作為水產(chǎn)養(yǎng)殖飼料的主要基礎(chǔ)原料,含有豐富的抗生素及ARGs,其對沉積物中ARGs豐度的影響有明顯劑量效應,其中的營養(yǎng)物質(zhì)可以驅(qū)動微生物群落結(jié)構(gòu)來改變沉積物中ARGs[69].重要的,全球魚粉產(chǎn)量近70%被用于水產(chǎn)養(yǎng)殖,而中國是全球魚粉的主要消費國[70],“飼料禁抗”是大勢所趨.

但是水產(chǎn)養(yǎng)殖環(huán)境中養(yǎng)殖用藥和飼料中抗生素只是ARGs發(fā)生和傳播的主要應激源,抗生素與其相應的ARGs并非總是同時出現(xiàn),單一類型的抗生素使用下,也可能存在復雜的ARGs污染[71-73]. ARGs也不是僅由于同源抗生素而引起.實際養(yǎng)殖過程中,盡管并未添加的抗生素,其ARGs照樣可以存在.如氯霉素、紅霉素類藥品早就被禁止用于水產(chǎn)養(yǎng)殖業(yè),但水東灣蝦場中卻是蝦塘水中主要ARGs,在數(shù)地被檢出,也均以一定量存在[44-46,48].

2.2 養(yǎng)殖模式對抗生素抗性基因的影響

養(yǎng)殖模式是水產(chǎn)養(yǎng)殖獨有的ARGs影響因素.不同單一養(yǎng)殖模式下,ARGs的多樣性及豐度皆有差異.Gu等[55]對比了瀛湖圍欄養(yǎng)殖和箱型養(yǎng)魚ARGs的污染情況,發(fā)現(xiàn)箱型養(yǎng)殖的ARGs多樣性要比圍欄養(yǎng)殖更高,其認為由于不同魚類養(yǎng)殖方式,理化參數(shù)變化不同,從而引起的微生物群落變化導致,通過調(diào)節(jié)湖泊的某些因素是可以控制ARGs的.而相比于單一養(yǎng)殖模式,在當前循環(huán)資源的多營養(yǎng)級養(yǎng)殖發(fā)展趨勢下,混合養(yǎng)殖(鴨-魚池)水體相比于單一養(yǎng)殖更不易受到ARGs污染[49].Yuan等[41]也發(fā)現(xiàn)單一牛蛙養(yǎng)殖場的沉積物中ARGs的檢測頻率和豐度均高于混養(yǎng)養(yǎng)殖場沉積物.但是,混合養(yǎng)殖對ARGs的影響還取決于混養(yǎng)品種.杭州灣河口養(yǎng)殖區(qū)報道[46]在混養(yǎng)池塘中,與雜食性魚類相結(jié)合的蝦養(yǎng)殖可能會增加ARGs豐度,而與濾食性魚類養(yǎng)殖可能會減少ARGs豐度.合理安排混合養(yǎng)殖模式以及養(yǎng)殖品種是控制ARGs的重要手段.

2.3 環(huán)境理化因子對抗生素抗性基因影響

水產(chǎn)養(yǎng)殖環(huán)境的理化因子包括水溫、鹽度、酸堿性、化學耗氧量、溶解氧、營養(yǎng)物質(zhì)和重金屬等.這些因子復雜多變,同氣候條件、放養(yǎng)密度、餌料種類和投餌量、生物排泄物等有密切關(guān)系.而針對水產(chǎn)養(yǎng)殖中環(huán)境理化因子對ARGs賦存狀況和傳播規(guī)律的影響的研究極少.養(yǎng)殖過程中某些環(huán)境理化因子是可控的,弄清其在水產(chǎn)養(yǎng)殖環(huán)境中影響,加以有效人為調(diào)控,可能達到控制ARGs污染的目的.

2.3.1 溫度、鹽度等對抗生素抗性基因影響 對于溫度,適宜的溫度更有助于水產(chǎn)品的生長.溫度也會影響質(zhì)粒的轉(zhuǎn)移過程,37℃下抗性質(zhì)粒更容易發(fā)生水平轉(zhuǎn)移[74].長江口及鄰近海域的研究[75]發(fā)現(xiàn)溫度與1和等基因的豐度之間呈顯著相關(guān)性(P<0.05).對于鹽度,其可調(diào)控微生物群落的組成和結(jié)構(gòu),從而調(diào)控環(huán)境中ARGs.河口開放養(yǎng)殖區(qū)最易受其影響,隨著鹽度的增加,垂直傳播的動態(tài)平衡會被打破,磺胺類ARGs會減少;鹽度影響的還具有季節(jié)性,不同季節(jié),鹽度對于ARGs有不同的影響,所影響的ARGs類型也有所不同[76].

總有機碳(TOC)、化學需氧量(COD)、總磷(TP)、總氮(TN)等水質(zhì)指標,代表著養(yǎng)殖環(huán)境的污染狀況,其皆影響ARGs.水東灣蝦塘水中和TOC呈正相關(guān)(<0.05);和TOC呈極強正相關(guān)(<0.01);與COD極強相關(guān)(<0.01)[48].東江流域[18]、長江口[77]的研究也表明TS、TOC、TN和等ARGs呈極顯著相關(guān)(<0.01).懸浮顆粒物(SS)還被認為可能是ARGs的載體,其豐度與、的豐度均存在顯著相關(guān)性(<0.05)[48,75].但在水產(chǎn)養(yǎng)殖過程中,TOC、COD、TP、TN等從機制上如何影響ARGs,還有待研究.適宜的含量既可保證水產(chǎn)品生長,水質(zhì)良好,又能降低ARGs的威脅.

至于酸堿性,養(yǎng)殖環(huán)境中pH值一般變化幅度較小,但不同ARGs受pH值變化影響有不同的機制.如Huang等[78]發(fā)現(xiàn)厭氧處理過程中酸性條件下,噬菌體DNA含量會增加,且更有利于eDNA的排泄,因此四環(huán)素ARGs水平轉(zhuǎn)移的數(shù)量和其持基因能力大大提高;而堿性條件下,接合轉(zhuǎn)移可能受到限制;Lin等[34]研究卻發(fā)現(xiàn)糞肥中,酸化降低了磺胺類ARGs的水平轉(zhuǎn)移,pH可誘導厚壁菌門的富集,增強磺胺類抗生素降解,從而抑制的水平基因轉(zhuǎn)移.

2.3.2 重金屬對抗生素抗性基因影響 重金屬對水產(chǎn)養(yǎng)殖環(huán)境中ARGs賦存及傳播有極大影響,部分微量金屬元素也是水產(chǎn)品生長所必須的因子之一[79].螯合銅、硫酸銅等重金屬鹽作為消毒劑也在水產(chǎn)養(yǎng)殖中廣泛應用[80].研究發(fā)現(xiàn)重金屬可與抗生素共同對微生物造成選擇壓力,且重金屬可以抵抗降解,選擇壓力可以長期施加,在同一質(zhì)粒中重金屬甚至可和ARGs共同存在[31,71,81].江蘇省白馬湖水產(chǎn)養(yǎng)殖場[52]評估Cu2+與ARGs呈明顯的正相關(guān)關(guān)系;東江流域[18]大部分ARGs與Cu、Zn等重金屬之間存在顯著的相關(guān)性.其中,弱酸提取態(tài)的重金屬被認為更容易從環(huán)境中釋放而被微生物利用.弱酸提取態(tài)Zn的含量與ARGs之間的顯著相關(guān)性更強(<0.01), Zn離子被認為是調(diào)控ARGs賦存的重要金屬元素之一[8,75].在機制上,已有銀、鋅影響機制的報道,但并不針對水產(chǎn)養(yǎng)殖環(huán)境.據(jù)報道,AgNPs和Ag+是通過誘導ROS過量產(chǎn)生,觸發(fā)SOS反應,增加細胞膜通透性來促進接合基因轉(zhuǎn)移,從而加強ARGs的傳播[82];天然閃鋅礦和ZnS納米顆粒則是促進細菌之間質(zhì)粒介導的ARGs水平轉(zhuǎn)移,增強抗性編碼質(zhì)粒的接合轉(zhuǎn)移[83].這從機制上解釋了重金屬對ARGs傳播的影響,為在水產(chǎn)養(yǎng)殖環(huán)境中進行類似研究提供了參考.而實際養(yǎng)殖過程中,重金屬在水產(chǎn)養(yǎng)殖環(huán)境中易富集、有毒性、去除難度高,除了對ARGs產(chǎn)生影響,對水產(chǎn)品和整個水產(chǎn)養(yǎng)殖環(huán)境也產(chǎn)生嚴重危害.探究重金屬對水產(chǎn)養(yǎng)殖環(huán)境中ARGs賦存和傳播的影響具有重要意義.

3 結(jié)語

為了從根本上防止抗生素耐藥性的發(fā)生和傳播,未來水產(chǎn)養(yǎng)殖環(huán)境中ARGs的研究可抓住以下幾點:

1) 深入解析ARGs在水產(chǎn)養(yǎng)殖環(huán)境、水產(chǎn)品以及人體致病菌之間的關(guān)系.盡管目前已發(fā)現(xiàn)水產(chǎn)養(yǎng)殖中的ARGs能通過食物鏈對人類造成威脅,和人類腸道菌群、致病菌所攜帶ARGs具有相似性,但缺乏更多證據(jù)及相關(guān)機制的研究.

2) 合理選擇水產(chǎn)養(yǎng)殖環(huán)境中代表性污染ARGs,開發(fā)適合水產(chǎn)養(yǎng)殖環(huán)境準確、快捷的ARGs監(jiān)測方法,系統(tǒng)的獲取水產(chǎn)養(yǎng)殖環(huán)境中ARGs的相關(guān)數(shù)據(jù),制定針對水產(chǎn)養(yǎng)殖環(huán)境ARGs污染的標準.傳統(tǒng)PCR檢測ARGs具有局限性,而宏基因組和高通量PCR在水產(chǎn)養(yǎng)殖環(huán)境的研究中相對較少,且檢測成本高.選定代表性的ARGs優(yōu)先控制,將ARGs污染監(jiān)測常規(guī)化,是控制ARGs污染的必要手段

3) 探究水產(chǎn)養(yǎng)殖過程中主流養(yǎng)殖模式、溫度、重金屬鹽消毒劑等可調(diào)因素對ARGs發(fā)生和傳播的影響,了解其影響機制,有效的把控養(yǎng)殖過程中ARGs的發(fā)生、傳播乃至去除.

4) 建立ARGs生態(tài)風險評估方法及預警體系,從而對水產(chǎn)養(yǎng)殖環(huán)境中ARGs對人類健康和生態(tài)環(huán)境的威脅科學的評估、預防ARGs污染的發(fā)生.

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Research progress on the occurrence and influencing factors of antibiotic resistance genes in aquaculture environment.

LI Shi-sheng1,2, GAO Hui2**, ZHAO Fu-qiang2,4, CHEN Yi-qin2, ZHANG Ke-yu1,2, HUANG Jia-jin1,2, NA Guang-shui1,2,3*

(1.College of Marine Ecology and Environment, Shanghai Ocean University, Shanghai 201306, China；2.National Marine Environmental Monitoring Center, Dalian 116023, China；3.College of Ecology and Environment, Hainan Tropical Ocean University, Sanya 572022, China；4.College of Marine Technology and Environment, Dalian Ocean University, Dalian 116023, China)., 2021,41(11)：5314～5325

It is important to clarify the contamination status of antibiotic resistance genes (ARGs) in the aquaculture environment in China.In this paper, the current status of the occurrence of seven major classes of resistance genes in aquaculture environmental water, sediment and aquatic products was counted. The factors influencing their occurrence were further analyzed. The future research on the pollution of ARGs in aquaculture in China was prospected. ARGs in the aquaculture environment are mainly in the form of iDNA.Sulfonamide and tetracycline resistance genes are considered to be the major ARGs in aquaculture environments due to their high abundance and detection rates, while the abundance and detection rate of quinolones, chloramphenicol, β-lactams, macrolides and aminoglycosides resistance genes are reported to be relatively low. In the process of aquaculture, aquaculture drugs, feed and culture mode are the main factors affecting the occurrence of ARGs in aquaculture environment, but the role of environmental physical and chemical factors in regulating ARGs in aquaculture environment cannot be ignored.

antibiotic resistance genes；aquaculture environment；occurrence state；influencing factors

X52

A

1000-6923(2021)11-5314-12

李十盛(1997-),男,吉林白山人,上海海洋大學碩士研究生,主要從事抗生素及抗生素抗性基因研究.

2021-04-07

國家重點研發(fā)計劃項目(2019YFD0901104);國家自然科學基金資助項目(41976222,42006195);自然資源部海洋大氣化學與全球變化重點實驗室開放基金(GCMAC2010)

* 責任作者, 研究員, gsna@nmemc.org.cn; **工程師, hgao@nmemc. org.cn