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汉坦病毒-copy from somewhere…

引言

汉坦病毒属于布尼亚病毒科的一个包膜病毒属，即汉坦病毒属(Orthohantavirus)。医学上重要的汉坦病毒均由鼠科与仓鼠科啮齿动物携带。这些病原体可导致两种重症急性发热性疾病：

●肾综合征出血热(hemorrhagic fever with renal syndrome, HFRS；由旧世界汉坦病毒引发)

●汉坦病毒心肺综合征(hantavirus cardiopulmonary syndrome, HCPS；由新世界汉坦病毒引发)，亦称汉坦病毒肺综合征(hantavirus pulmonary syndrome, HPS)

本文将总结汉坦病毒感染(特别是HCPS)的流行病学与诊断。汉坦病毒感染的临床表现、发病机制和预防见其他专题。(参见 “汉坦病毒心肺综合征”和 “汉坦病毒感染引起的肾脏受累”和 “汉坦病毒感染的发病机制”)

病毒学

目前确切识别出了多少种汉坦病毒尚有争议，但自然界中至少有30种不同的汉坦病毒，至少十多种可导致人类疾病 (表 1)。首先在啮齿动物身上发现，后来被证明是人类病原体的汉坦病毒例子很少[1,2]。

汉坦病毒具有单链反义RNA基因组，可分为3个片段。L片段(或大片段)编码复制酶，即RNA依赖性RNA聚合酶与核酸内切酶；M片段(中片段)编码包膜糖蛋白Gn和Gc；S片段编码核衣壳蛋白N[3,4]。

包膜糖蛋白可通过全身的内皮细胞和血小板的细胞表面整合素-β3介导汉坦病毒附着于细胞[3,5]。最近还发现了其他潜在的细胞侵入相关分子[6]。汉坦病毒在体内也会感染血管内皮细胞。动物模型和人体研究证实，某些汉坦病毒对Kupffer细胞与肾上皮细胞具有更广泛的趋向性。

啮齿动物宿主

虽然已知各汉坦病毒能感染不同种类的哺乳动物(包括啮齿动物、食虫动物、蝙蝠)，但每种致病性汉坦病毒都与特定野生啮齿动物相关，后者为其主要自然宿主[7,8]。

在携带汉坦病毒的主要啮齿动物中，一般5%-20%有该病毒的抗体，而且血清反应阳性的动物通常有持续的活动性感染。汉坦病毒分布在全球各地，与其各自的宿主栖息地一致。

●引起HFRS的汉坦病毒由田鼠携带，但普马拉病毒除外，它由棕背䶄携带。

●北美与南美的许多本土啮齿动物携带有HCPS病原体。鹿鼠为辛诺柏病毒(Sin Nombre Virus, SNV)的主要宿主，SNV是北美地区HCPS的最主要病因，但公认存在其他啮齿类宿主的溢出感染。

HCPS病原体之间的关联比它们与HFRS病原体之间的联系更密切[3,9]。关系密切的病毒感染亲缘关系密切的宿主并且引起相似的临床综合征，这一发现使人们认为，数千年来汉坦病毒属在其主要宿主种内共同进化[3]。然而，有假说认为，汉坦病毒在同一地区的啮齿类动物中水平传播是其演化和多样化的一个重要驱动因素[10]。

在鱼类和爬行动物等物种中也发现了汉坦病毒[11]。据报道，曾有血清学证据显示人类暴露于来自非啮齿动物宿主的汉坦病毒，但尚未发现特定疾病与这种暴露有关[12]。

传播

接触啮齿动物是汉坦病毒传播到人类的一个重要因素。许多汉坦病毒由急性感染的宿主啮齿动物通过尿液、粪便或唾液排出。研究者怀疑，汉坦病毒传播给人类主要是通过气溶胶途径，尽管气溶胶的确切性质难以确定(主要是由于该病毒感染罕见)。

极少数感染汉坦病毒的患者自述近期被啮齿动物咬伤。然而，许多患者自述在出现症状前2-4周遇到过活的或死的啮齿动物，或者进入过有明显啮齿动物出没证据的房间。室内暴露可能尤其重要，尤其是当前啮齿动物出没较活跃时。例如，在美国几乎所有辛诺柏汉坦病毒感染者都表示曾进入有啮齿动物出没的建筑物[13]。血清流行病学研究表明，北美啮齿动物职业性接触者感染汉坦病毒属的风险较低[14,15]。

欧洲、亚洲(尤其是俄罗斯)对HFRS实验室获得性感染病例进行了研究。在1961年俄罗斯的暴发中，113例病例与在通风不良、维护不当的动物研究机构中饲养感染汉坦病毒的啮齿动物有关。许多实验室相关病例与啮齿动物最为密切的接触为短时间进入饲养已感染宿主啮齿动物的实验室。大量此类病例支持经空气传播是人类暴露的常规途径[16]。现代的实验室操作规范使得已经很难新发生来自实验室的汉坦病毒大规模暴发。

汉坦病毒在人与人之间的传播较为罕见，且仅限于一个汉坦病毒种，即安第斯病毒。有时难以区分是家庭成员之间的传播还是啮齿动物的同源暴露[17,18]。汉坦病毒感染的暴露与症状出现之间的时间间隔为1-6周，中位时间为14-17日。初始病例发病7日内发生的继发病例通常被归为同源暴露，而非人传人[18]。

智利的一项前瞻性研究纳入了76例安第斯病毒感染患者的476例家庭接触者，指示病例的性伴侣罹患HCPS的风险为17.6%，而其他家庭接触者为1.2%[19]。在症状发作或特异性抗体出现前5-15日，通过逆转录酶聚合酶链反应(reverse-transcriptase polymerase chain reaction, RT-PCR)，在外周血单个核细胞(peripheral blood mononuclear cell, PBMC)中可检出病毒RNA。在普马拉病毒所致HFRS患者的唾液中发现了汉坦病毒RNA，这提示病毒可能经口腔分泌物传播，但唾液中未检出活病毒，而且流行病学研究结果不支持该传播途径[20]。

有研究者尝试确定寄生螨是否参与汉坦病毒在自然界中的携带或传播[21,22]，但支持证据仅为推测性[22]。

汉坦病毒的发现

几个世纪前很可能就有过汉坦病毒暴发。在西方医学发现汉坦病毒病后，回顾中国古籍发现，大约公元960年就记载了符合此病的综合征[23]。

1951-1953年朝鲜战争期间联合国军队暴发了HFRS，研究者进行了详尽描述，引起了西方医学界的关注。2000多名士兵患病，病死率约为5%[24]。研究者怀疑该疾病的流行病学可能涉及啮齿动物，但20多年后才从黑线姬鼠体内分离出病原体，即汉滩病毒[25]。在接下来的几年，韩国(汉城病毒)和欧洲(普马拉病毒与多布拉伐病毒)发现了HFRS的其他病原体[26-28]。

在啮齿动物群体中对汉坦病毒暴露的血清学调查表明，汉坦病毒在美国分布广泛[29]。然而，一直在新世界搜寻HFRS的研究人员却意外发现了HCPS。20世纪80年代，在美国分别从共生鼠[褐家鼠(Rattus norvegicus)]和一种棕背䶄中分离出汉城病毒和一种新汉坦病毒种(希望山病毒)[30,31]。经过长达10年的研究，在美国发现了极少数由汉城病毒引起的HFRS人类病例[32]。此类病例似乎非常罕见[33,34]，但2016年出现过一次暴发。(参见下文‘暴发’)

1993年春季，在新墨西哥州与亚利桑那州之间北部边界处的纳瓦霍部落成员中发现了较多的原因不明的发热及急性呼吸窘迫综合征(acute respiratory distress syndrome, ARDS)病例[35]。这一最初发病的患者人群中病死率约为80%。这些患者的血清样本中含有可抗来自韩国和芬兰的汉坦病毒分离株的IgM抗体。在后来的数周，证实了新发现的SNV为HCPS的病原体[36-40]。研究人员后来发现了其他新型汉坦病毒，它们是美国HCPS罕见病例的病原体和南美洲和中美洲HCPS常见病原体[41-46]。自1993年的暴发以来，美国已有41个州发现了病例，大多非印第安人[47]。

全球患病率

许多汉坦病毒病的病例很可能未上报，部分原因是临床上将其与钩端螺旋体病、登革热、疟疾、COVID-19及其他疾病相混淆，尤其是在热带地区。目前，中国的汉坦病毒病年发病率最高。中国每年报告的HFRS病例有16,000-100,000例或更多，俄罗斯每年可能出现数千例 (表 1)。中国的HFRS病例目前最常由汉城病毒引起，有些由汉滩病毒引起。

在整个欧洲，每年由普马拉和多布拉伐病毒引起的HFRS为数千例，但确切数字未知。流行性肾病(普马拉病毒–HFRS)的病死率<1%，而汉滩病毒或多布拉伐病毒HFRS为2%-10%[3]。中国的病死率为1%，其中女性患者的病死率略高于男性[48]。2012年，在英国发现2例由汉城病毒引起的HFRS；其中一例溯源至野鼠，而另一例为宠物鼠[49,50]。在美国也有极少数由汉城病毒引起的HFRS[32-34]，但2016年发生过一次汉城病毒感染暴发。(参见上文‘汉坦病毒的发现’和‘暴发’)

截至2021年12月，美国共确诊了850例HCPS，包括1993年发现该病后回顾确定的一些病例[51-57]。在美国，1993-1994年、1999-2000年、2006年1-3月报道的HCPS病例数增多[58]。这些增长与先前发生的厄尔尼诺南方振荡(El Niño Southern Oscillation, ENSO)气候事件有时间关联，但也可能是巧合[13]。

每年10%-15%的北美病例来自加拿大[59]。其他出现过汉坦病毒的国家包括：阿根廷、玻利维亚、巴西、智利、厄瓜多尔、巴拿马、巴拉圭、秘鲁、乌拉圭、法属圭亚那和委内瑞拉[2,60-62]。整个美洲共诊断了3000多例HCPS，远低于欧亚HFRS的总病例数。然而，HCPS的病死率通常为25%-50%，远高于HFRS。HCPS治疗通常效果欠佳，而利巴韦林可能对汉滩病毒所致HFRS有效。(参见 “汉坦病毒心肺综合征”)

所有受汉坦病毒病影响的国家中，一些地区的感染发病率较高，而其他地区极少。美国西部和西南部各州的病例远多于中西部或东部各州，如新墨西哥州、亚利桑那州、加利福尼亚州、华盛顿州、科罗拉多州、蒙大拿州以及德克萨斯州[56]。美国西部各州(如加利福尼亚州、新墨西哥州和科罗拉多州)的山地生物群落是对鹿鼠尤其有利的环境，这些地区的人类感染往往发生在海拔2000m以上[63]。由ENSO等造成的气候影响可能是发生汉坦病毒病的重要因素，但ENSO影响HCPS发生的机制仍有待进一步研究[13]。汉坦病毒病的高发地区变化无常，气候变化的潜在影响也很难预测。

暴发

正如上文所述，1951-1953年朝鲜战争期间联合国军中发生的一次大规模暴发使西方医学界注意到HFRS。(参见上文‘汉坦病毒的发现’)

后来又有一些由汉坦病毒引起的暴发：

●1993年春季，在新墨西哥州与亚利桑那州之间北部边界处的纳瓦霍部落成员中，发现了较多不明原因发热伴ARDS病例，最后确认病因为SNV。(参见上文‘汉坦病毒的发现’)

●2012年夏季，在加利福尼亚州约塞米蒂国家公园的10名游客中发生了汉坦病毒感染暴发，9例为HCPS，3例死亡[64,65]。9例患者曾宿于一组孤立的帐篷屋中。在孤立区发现了啮齿动物出没。

●2016年12月，美国田纳西州一名饲养宠物鼠的女性出现了急性发热性疾病，伴血尿和血清肌酐升高，检查发现其感染了汉城病毒[66]。她后来康复，与该指示患者同住的一名家庭成员在处理鼠类排泄物后也发生疾病且检测结果为阳性。还有一例于2018年报道自哥伦比亚特区[67]。

●2017年1月，美国威斯康星州鼠类饲养员和宠物鼠主人中暴发汉城病毒感染[66,68,69]。后续调查发现这次暴发中有24人感染了汉城病毒，其中涉及美国11个州和加拿大的鼠类养殖场、宠物商店和宠物鼠饲养家庭[69]。关于潜在暴露个体的检测推荐以及其他信息可参见美国CDC网站(United States Centers for Disease Control and Prevention's website)。

欧洲人感染汉城病毒通常是因为接触了宠物鼠[70]。英国的商业繁殖基地中也发现了感染的鼠类，目前可能需要进一步干预以降低全世界潜在啮齿类动物拥有者的感染风险[71]。

血清阳性率

人类既往暴露于汉坦病毒(根据特异性IgG抗体阳性确定)的情况在世界各地差异很大。关于无症状感染发生率的流行病学数据不一致。一项基于人群的研究确定汉滩病毒有症状感染与无症状感染的比率约为1:5[72]。而其他地区有症状感染占感染总数的比例相当低。巴拉圭查科土著人群虽然有40%的血清阳性率，但临床HCPS发病罕见[73,74]。巴拿马也报道了相似的高血清阳性率[75]。汉坦病毒抗体相对持久，有病例在出现临床症状约20年甚至长达36年后仍可根据血清特异性IgG轻松得以诊断[53-55]。

尚不清楚是宿主间的差异还是特定汉坦病毒间的毒力差异造成了感染率和发病的差异。在美国，大多数受累人群居住在亚利桑那州、科罗拉多州、新墨西哥州和犹他州，这些地区的血清阳性率约为1%[76-78]。但美国成人的血清阳性率可能低于0.1%。儿童很少患HFRS或HCPS，但南美洲除外，南美洲曾有年仅5岁的儿童死于HCPS[79]。然而，2009年5例6-14岁的美国西部儿童发生HCPS[78]。美国后来数年内又出现了几例病例，其中包括年龄低至5岁的儿童[80]。

血清阳性率在密切接触鼠类的人群中很可能更高，但可能因鼠的类型和暴露程度而异。在英国的一项血清学阳性率研究中，大约1/3的宠物花枝鼠(一个常见驯养鼠品种)主人和饲养员有汉城病毒抗体，而职业性暴露于宠物鼠或野生鼠人群的血清阳性率并不高于一般人群[81]。这些数据表明，宠物花枝鼠的拥有者和饲养员发生临床或亚临床汉坦病毒感染的风险尤其高。

汉坦病毒活动的季节性和周期性

汉坦病毒感染呈季节性暴发，且不同年份和地区发病率差异较大。

●在中国，HFRS暴发出现于春耕季节，而秋季更甚。农民参与秋收时可能因为睡在有啮齿动物出没的非密闭小屋内而接触汉坦病毒。

●在斯堪的纳维亚，HFRS每几年就会周期性暴发一次，而且似乎之前均有野生棕背䶄大量繁殖[82-84]。

●2001年智利的人类感染暴发与安第斯病毒的鼠类宿主长尾小啸鼠(Oligoryzomys longicaudatus)的突然增多有关[85]。

感染模式

美国和加拿大的HCPS病例为散发。

而在智利，25%-30%的HCPS病例为聚集性[86]。通常认为这些聚集出现的患者是暴露于共同感染源，例如同一只活跃排出异常大量病毒的感染性啮齿动物。然而，一些患者在指示病例出现症状后多日或数周才发生症状，且距离啮齿动物出没地较远，怀疑为人传人[19]。

已有妊娠期出现HCPS的病例。一项病例系列研究显示，与历史对照HCPS患者(非妊娠)相比，5例妊娠患者的临床表现并无差异[87]。报告了1例产妇死亡与2例胎儿死亡，但未发现SNV垂直传播的证据。母乳的SNV RNA检测可呈阳性，但尚未确定母乳喂养与病毒传播有关联[88]。新生儿汉坦病毒感染极其罕见[89]。根据世界范围内的经验，妊娠期发生汉坦病毒感染时，母胎/婴预后较差，但尚未报道该病毒引发的具体病理改变(像风疹、寨卡病毒那样)[90]。

诊断

在大多数实验室中，血清学方法是诊断人类或啮齿动物汉坦病毒感染的首选方法。虽然已制定了流程图以帮助临床鉴别汉坦病毒感染与钩端螺旋体病，但还不足以确诊[91]。

血清学检查 — 血清学检查是诊断急性或既往汉坦病毒感染的主要方法，不过均未获得美国FDA的批准。到症状明显时，患者体内均已出现IgM类抗病毒抗体，而且大多数会有IgG类抗体[92,93]。诊断性检测包括：酶联免疫吸附试验(enzyme-linked immunosorbent assay, ELISA)、条带免疫印迹试验(strip immunoblot tests, SIA)、Western印迹试验、间接免疫荧光法(indirect immunofluorescence, IFA)、补体结合试验、血凝抑制试验，以及灶点或蚀斑减少中和试验，用以检测汉坦病毒抗体[94]。

急性感染时存在特异性抗汉坦病毒IgM(通常应用核衣壳或N抗原)或抗汉坦病毒IgG滴度可增至4倍，由此可与既往感染相鉴别。

在美国，提供汉坦病毒感染诊断性检测的州立卫生部门使用美国CDC研制并分发的IgG检测和μ链-IgM捕获法ELISA。这些ELISA采用重组表达的N抗原。加拿大也已采用此项检查[93]。也有人研发了一项蛋白质印迹检测，利用重组抗原以及区分IgM与IgG的同型特异性复合物，其检出率与前法类似。

诊治HFRS的亚洲医院常采用磁珠凝集试验(HantaDia)、ELISA以及IFA方法诊断汉滩病毒与汉城病毒感染[95]。欧洲实验室多采用ELISA，有部分实验室采用通过重组DNA技术合成的蛋白抗原，但仍有一些实验室采用IFA[96-99]。南美洲参考实验室通常采用ELISA[100]。

聚合酶链反应 — 对于可引起高病死率暴发性感染的汉坦病毒(安第斯病毒、辛诺柏病毒、多布拉伐病毒，还可能包括汉滩病毒[101])，在肺病早期采用PBMC或血清进行巢式RT-PCR一般可检测到病毒RNA。然而，病毒RNA通常在数日后从循环中消失，因此血清学技术的诊断准确性较高。普马拉病毒通常只引起轻度感染，仅可在少数患者中检测到RNA，通常为感染异常严重的患者。因此，只有极少数的临床实验室提供常规RT-PCR诊断性检查。RT-PCR更适合作为一种备用检测或确诊性检测，或者用于识别可能的感染部位[94]。

尸检 — 尸检时，使用针对病毒N抗原的抗体通过免疫组化方法很容易检测出病毒抗原。N抗原染色在细胞质中点状分布，尤其集中在肺的血管内皮以及肾的肾小球毛细血管内皮[102,103]。可应用巢式RT-PCR检测尸检采集的冰冻切片或固定的石蜡包埋组织中的病毒RNA[37,40]。

分子流行病学

对单个病毒遗传序列的检测已用作分子流行病学研究工具。虽然特定病毒基因型在一个小范围地区可多年保持稳定，但几乎不可能在几千米外找到另一种序列相同的病毒[104]。因此，用人体样本进行RT-PCR扩增后，对PCR产物进行DNA测序，可提供相关病毒种及准确暴露位置的有用信息。通过比较特定患者与附近受感染啮齿动物或同一区域内其他患者的病毒RNA序列，也许能区分家庭周围与工作场所获得的感染。

为了进行此类检测，有必要在各个可能的暴露场所收集至少3-10只血清反应呈阳性的啮齿动物。根据新墨西哥大学感染病与免疫中心的经验，在经过充分调查的地方，在3次尝试中至少有2次可鉴定出序列与患者的病毒序列相同的病毒[104]。

总结

●啮齿动物宿主–旧世界[肾综合征出血热(HFRS)]和新世界[汉坦病毒心肺综合征(HCPS)]汉坦病毒均由野生啮齿动物携带和传播。感染以地方性流行和其他流行形式出现，通常是天然啮齿动物大量繁殖的结果。全世界已鉴定出11种汉坦病毒病原体，其分布与其啮齿动物携带者一致 (表 1)。患者通过啮齿动物排泄物的气溶胶感染病毒，一般在通风不良的封闭建筑中或打扫时发生。(参见上文‘引言’和‘啮齿动物宿主’和‘传播’)

●地理分布–中国的汉坦病毒疾病年发病率最高。中国每年报道的HFRS病例为16,000-100,000例或更多，俄罗斯每年可能出现数千例。相比之下，美国仅确诊了833例HCPS。世界其他地方也有汉坦病毒病的病例，如欧洲与南美洲 (表 1)。(参见上文‘全球患病率’)

●死亡率因病毒种而异–病死率很大程度上取决于感染的病毒种。在世界上一些地区，有证据表明存在相当多无症状或亚临床感染，但在其他一些地区难以找到已暴露于病毒而未发病的个体。(参见上文‘汉坦病毒的发现’和‘全球患病率’)

●血清学诊断–急性和既往汉坦病毒感染的主要诊断方法均是血清学检查。在症状明显时，患者体内均已出现IgM类抗病毒抗体，而且大多数会有IgG类抗体。急性感染时存在特异性抗汉坦病毒IgM(通常应用核衣壳或N抗原)或抗汉坦病毒IgG滴度可增至4倍，由此可与既往感染相鉴别。(参见上文‘血清学检查’)

●PCR的作用–对于引起高病死率暴发性感染的汉坦病毒(安第斯病毒、辛诺柏病毒、多布拉伐病毒，也可能包括汉滩病毒)，在肺病早期采用外周血单个核细胞(PBMC)或血清进行巢式逆转录酶聚合酶链反应(RT-PCR)一般可检出病毒RNA。然而，病毒RNA通常在数日后从循环中消失。(参见上文‘聚合酶链反应’)

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