摘要
HFRS疫源地性质类型的空间异质性具有空间尺度依赖性,空间尺度不同其空间异质性特征不同。由于其以往研究所采用的空间尺度不够细化,不能搞清微小生态环境内的HFRS病原体与宿主鼠类间的时空流行病学和生态学联系,因而不能有效的指导HFRS疫点范围的划定以及疫点内采取以灭鼠与疫苗接种为主的预防干预措施。为此,本研究从多学科交叉入手,将空间流行病学、群落生态学、分子生物学、分子遗传学、空间信息统计学等学科的理论方法有机结合,以GIS为空间数据管理分析平台,在微小尺度上(以户为空间尺度单位),圈定山东省莒南县两个试验疫区(约6×6km),采用纵向监测方法,于2006年2月-2007年1月在试验疫区进行HFRS宿主动物和病原学监测。深入分析试验疫区内宿主动物及其寄生螨类的种群结构和生态位特征,以阐明试验疫区内HFRS病毒与宿主鼠类之间的时空动态关系。
(一)主要研究结果:
1、HFRS的宿主鼠类生态特征
宿主动物的群落结构因地理生态环境的不同具有一定差异,试验疫区各自然村内以褐家鼠和小家鼠为优势鼠种,二者在生态位占据明显优势地位。宿主鼠类的生态学特征表现为:①不同试验疫区内宿主鼠类的群落存在一定差异;②不同自然村内的宿主鼠种群落结构的季节分布差异较大:③小家鼠和褐家鼠的空间、时间生态位宽度均很高,褐家鼠和小家鼠的空间、时间生态位重叠指数最大:④由于自然条件、地理因素的影响,两试验疫区的宿主鼠类群落多样性指数、生态位重叠指数存在一定的差异,宿主鼠生态状况存在不同;⑤各鼠种之间的食物生态位重叠指数均很高;各鼠种之间的生殖生态位重叠指数均很高。
2、小地域生态环境内宿主鼠类分布特征
从试验疫区整体看,两试验疫区第一优势鼠种均为小家鼠,第二优势鼠种均为褐家鼠。
(1)第一试验疫区,各自然村鼠密度基本相同,但它们的主要鼠种构成却各有特点,西兰敦、东兰敦、曲流河和大山前4个自然村的鼠种构成相似,均以褐家鼠为主,小家鼠次之,其中,西兰敦和东兰敦鼠种构成中黑线姬鼠也占有较大比重;后良店、东良店和尤庄子3个自然村的鼠种构成相似,均以小家鼠为主,褐家鼠次之,且其它鼠种所占比例很小。
(2)第二试验疫区内,各自然村鼠密度也相差不大,但普遍明显低于第一试验疫区,各自然村的鼠种构成也相似,均以褐家鼠及小家鼠为主,其中,洼子和中店头的主要鼠种构成以小家鼠为主,褐家鼠次之;西店头以褐家鼠为主,小家鼠次之,其它鼠种所占比例很小。
3、小地域范围内鼠密度的季节消长
试验疫区内鼠密度具有明显的季节性特征,春(3-5月)、秋初(11月)、冬(12-2月)季密度高,夏(6-8月)、秋(9-10月)季密度低。反映了鼠类迁徙的季节分布特征:6-10月份室内密度最低,此期间内鼠类由室内向野外迁移,自11月份随着气候变冷,野外食物短缺,鼠类向室内回迁,因此11月份至次年5月份室内鼠密度较高。
4、影响宿主鼠类滋生的生态因子
本研究采用负二项回归方法探讨以户为单位宿主鼠类的聚集性以及影响聚集性的生态因子。发现造成宿主鼠类聚集分布的主要影响因素有以户为单位实施经常性灭鼠、水泥防鼠地板、庭院内禽畜放养、粮仓防鼠措施、杂物衣物食物混放等。其中,居民户自家实施经常性灭鼠、设水泥防鼠地板和实施粮仓防鼠措施是降低鼠密度的有效因素,而庭院内禽畜放养和杂物衣物食物混放是增加鼠类滋生的主要因素。
5、宿主鼠类的寄生螨类分布特征
从整个试验疫区看,褐家鼠虽然在捕获数量上低于小家鼠,但感染螨类的褐家鼠数量上却明显高于小家鼠;寄生革螨数量明显高于其他螨类。对各自然村螨类寄生情况逐一进行研究表明,虽然各自然村情况略有不同,但得出的结果与试验疫区整体情况相似:革螨是试验疫区宿主动物的优势寄生螨类,褐家鼠是螨类主要寄宿鼠类。
6、宿主鼠类带毒、感染特征
试验疫区室内主要宿主为褐家鼠、小家鼠,虽然小家鼠在数量上多于褐家鼠、但褐家鼠带毒率明显高于小家鼠。
(1)在一、二试验疫区内所捕获鼠类中均存在少量黑线姬鼠带毒的情况。
(2)在第一试验疫区内虽然鼠密度不高,但是鼠带毒率高,所以试验疫区内仍存在HFRS爆发的条件。
(3)试验疫区内宿主鼠类带毒率也呈明显的季节性波动,2月、5月、9～10月、12月的带毒率较高。
(4)褐家鼠和小家鼠不同性别、年龄间的带毒率、感染率均不存在差异。
7、宿主动物的空间流行病学特征
在GIS框架内,采用点格局分析方法,对全部捕获的宿主鼠类进行空间自相关聚集分析和空间“热点”分析。结果发现:
(1)在第一试验疫区内:①宿主鼠类的第一聚集区的平均半径为4.29米;最强聚集区的平均半径为14.43米;最大聚集区的平均半径为86.26米。各村聚集“热点”数差别较大,其一阶波动范围为3-8个,二阶波动范围为0-1个。②小家鼠的第一聚集区的平均半径为4.86米;最强聚集区的平均半径为21.14米;最大聚集区的平均半径为92.57米。各村聚集“热点”数差别较大,其一阶波动范围为1-12个,二阶波动范围为0-2个。③褐家鼠的第一聚集区的平均半径为5.00米;最强聚集区的平均半径为32.71米;最大聚集区的平均半径为93.86米。各村聚集“热点”数差别较大,其一阶波动范围为3-11个,无二阶聚集“热点”。
(2)在第二试验疫区内:①宿主鼠类的第一聚集区的平均半径为3.30米;最强聚集区的平均半径为13.67米;最大聚集区的平均半径为83.00米。各村聚集“热点”数差别较大,其一阶波动范围为3-7个,二阶波动范围为0-1个。②小家鼠的第一聚集区的平均半径为3.67米;最强聚集区的平均半径为21.00米;最大聚集区的平均半径为79.67米。各村聚集“热点”数差别较大,其一阶波动范围为2-5个,二阶波动范围为0-1个。③褐家鼠的第一聚集区的平均半径为3.67米;最强聚集区的平均半径为14.00米;最大聚集区的平均半径为68.33米。各村聚集“热点”数差别较小,其一阶波动范围为2-4个,无二阶聚集“热点”。
(3)最近邻空间系统聚类“热点”分析发现,各自然村中的宿主鼠类均存在明显的空间聚集“热点”。
8、病毒序列的分析揭示了宿主动物的活动轨迹
用系统进化树和最小生成树的方法描述分析了不同宿主鼠类所携带的SEO病毒株之间的空间关系,发现:
(1)各自然村之间的SEO病毒具有相对独立性。(2)各自然村间周边接壤地区宿主鼠类所携带的SEO病毒具有一定同质性。(3)HFRS病毒在试验疫区内呈现出一定的空间结构性。(4)同一自然村内的宿主鼠类所携带的病毒也呈现出一定的空间结构性和空间异质性。例如,在第一试验区大山前村内,部分宿主鼠类所携带病毒,构成明显的传播轨迹。
二、主要研究结论
1、试验疫区内的宿主鼠类的生态位特征
(1)宿主动物的群落结构因各自然村内的地理生态环境的不同而具有一定差异,但褐家鼠和小家鼠始终为各自然村内的优势鼠种。
(2)不同微生态环境内宿主鼠类的时间生态位、空间生态位、食物生态位、生殖生态位各有特点,提示在微小空间尺度上HFRS宿主动物的生态学种群结构仍十分复杂。
2、试验疫区内的宿主鼠类的分布特征及其影响因素
在微小生态环境内,宿主鼠类的鼠密度具有明显的季节变动性,空间分布呈现聚集特征,居民户自家实施经常性灭鼠、设水泥防鼠地板和实施粮仓防鼠措施是降低鼠密度的主要因素,而庭院内禽畜放养和杂物衣物食物混放是增加鼠类滋生的主要因素。
3、试验疫区内的宿主鼠类寄生螨类感染特征
革螨是试验疫区宿主动物的优势寄生螨类,褐家鼠是螨类主要寄生鼠类。
4、试验疫区内的宿主鼠类HFRS病毒感染特征
在试验疫区内,宿主鼠类的密度虽然很低,但其带毒率却很高,且褐家鼠带毒率明显高于小家鼠。虽然各自然村内的宿主鼠类的构成差别很大,但各自然村各鼠种之间带毒率的性别、年龄、季节分布特征基本一致。
5、试验疫区内的宿主鼠类空间流行病学特征
在微小的地域环境内,宿主鼠类最初聚集范围均与居民居室范围相当,最强的聚集区集中在1-2户居民的范围,最大聚集区在约80m的范围。试验疫区内各自然村的聚集“热点”数差别较大。
6、带病毒宿主动物的空间生态学和空间流行病学特征
(1)各自然村之间的SEO病毒间流行病学联系不明显。
(2)各自然村间周边接壤地区宿主鼠类所携带的SEO病毒具有一定同质性,提示鼠类可能由于在迁徙到野外的过程中相互接触感染了病毒,在向室内回迁的过程中又把病毒带回村内。
(3)HFRS病毒在试验疫区内呈现出一定的空间结构性,说明在自然村之间可能存在某些生态学屏障。
(4)同一自然村内的宿主鼠类所携带的病毒也呈现出一定的空间结构性和空间异质性。例如,在第一试验区大山前村内,部分宿主鼠类所携带病毒,构成明显的传播轨迹。说明在自然村之内既存在一定的生态学屏障,又具有一定的生态学联通轨迹。
Spatial heterogeneity of HFRS epidemic focus has the spatial criterion dependence. So the study of spatio-temporal dynamic of epidemics of HFRS should pay attention to spatial criterion.At present,there are many studies revealed spatio-temporal character of HFRS,including large spatial scale,middle spatial scale,Meso-micro scale,small scale.But spatio-temporal dynamics of HFRS virus and host in small area hasn't expounded by micro scale.The study on spatio-temporal dynamics of HFRS epidemics in small area has a great significance for working out practical prevention and control measures suited to the time and local conditions.For this reason,this study combines spatial epidemiology,community ecology,molecular biology,molecular genetics and spatial information statistics,based GIS.Longitudinal surveillance of host rats carried out in Jvnan County from Feb.2006 to Jan 2007,further expounding the relationship of spatio-temporal dynamic of HFRS various and host rats in small area.
Results:
1.Ecological characteristics of host rats
Ecological environment influences community structure of host rat in experimental district.Rattus norvegicus and Mus musculus are major species,either in time niche or spatial niche.
The ecological characteristics of host rat in experimental district as follow:
①The community structure of host mice is different in experimental districts.②The season distribution of community structure is different.③Both time niche breadth and spatial niche breadth of Rattus norvegicus and Mus musculus are very high.Both time niche overlap index and spatial niche overlap index of Rattus norvegicus and Mus musculus are highest.④Shannon-Weiner S indiex,Niche Overlap index and ecological status of host mice in two experimental districts are difference due to natural condition and geographical factor.⑤Food niche overlap index and breed niche overlap index between species are high.
2.The distribution character of host rat in small area
The frist major specie is Mus musculus,second major specie is Rattus norvegicus in experimental districts.
(1)Rat densities of seven villages in first experimental district are basically same.The constitutions of rat species in seven villages have different characteristics.The constitutions of rat species are basically similar in Xilandun village,Donglandun village,Qnliuhe village and Dashanqian village.The Rattus norvegicus is frist major specie,Mus musculus is second major specie in these four villages.The constitutions of rat species are basically similar in Houliangdian village,Dongliangdian village,Youzhuangzi village.The Mus musculus is frist major specie,Rattus norvegicus is second major specie in these three villages.
(2)Rat densities of three villages in second experimental district are basically same, lower than first experimental district.The constitutions of rat species in three villages are similar.The Rattus norvegicus is frist major specie,Mus musculus is second major specie in Xidiantou village.The Mus musculus is frist major specie,Rattus norvegicus is second major specie in Wazi village and Zhongliangdian village.
3.The season distribution of rat density in small area
The rat densities in Feb,March,April,May,Nov,and Dec are high.Other months are low.The season distribution of rat density reflects host rat migration.From June to Oct,host rats move to field.In winter,host rats back to inside.
4.Influence factors of host rats distribution
The spatial distribution shows clustered character.Regular deratization,cement anti-rat floor and methods used by residents are the main reasons for the decrease of rat density,while breeding livestock in yards and mixing up food,clothes and other goods are the main reasons for the increase of rats.
5.The distribution character of parasitic mite
In experimental district,the amount of Rattus norvegicus infected are more than Mus musculus infected.Gamasid mite is major mite in experiment.
6.The characteristics of the toxin-carrying rate of host rats
The major hosts in experiment districts are Rattus norvegicus and Mus norvegicus. Although the number of Mus norvegicus preponderates over that of Rattus norvegicus, the toxin-carrying rate of the former is much higher than that of the latter.
(1)In the captured rats in the first and the second experiment districts,there exist some toxin-carrying Apodemus.
(2)The toxin-carrying rate in the first experiment district is high,although the rat density there is low.Thus,the condition of the outburst of HFRS exist as well.
(3)The rat density and the toxin-carrying rate of host rat change according to the season.The toxin-carrying rates in February,May,September,October,and December are high.
(4)There exist no differences in toxin-carrying rate and infection rate between Rattus norvegicus and Mus norvegicus.
7.The spatial epidemiology characteristics of host animals
The spatial autocorrelation analysis and spatial "hot spot" analysis are performed by using point pattern analysis based on GIS.
(1)In the first experiment district:①The average radius of the first experiment district of host rat is 4.29 meters;The average radius of the strongest cluster district of host rat is 14.43 meters;The average radius of the biggest cluster district of host rat is 86.26 meters.The numbers of "hot spot" in different villages differ greatly.The undulation range of the first order is 3-8,and the undulation range of the second order is 0-1.②The average radius of the first experiment district of Mus norvegicus is 4.86 meters; The average radius of the strongest cluster district of Mus norvegicus is 21.14 meters; The average radius of the biggest cluster district of Mus norvegicus is 92.57 meters. The numbers of "hot spot" in different villages differ greatly.The undulation range of the first order is 1-12,and the undulation range of the second order is 0-2.③The average radius of the first experiment district of Rattus norvegicus is 5.00 meters;The average radius of the strongest cluster district of Rattus norvegicus is 32.71 meters; The average radius of the biggest cluster district of Rattus norvegicus is 93.86 meters. The numbers of "hot spot" in different villages differ greatly.The undulation range of the first order is 3-11,and there are no cluster hot spot of the second order.
(2)In the second experiment district:①The average radius of the first experiment district of host rat is 3.30 meters;The average radius of the strongest cluster district of host rat is 13.67 meters;The average radius of the biggest cluster district of host rat is 83.00 meters.The numbers of "hot spot" in different villages differ greatly.The undulation range of the first order is 3-7,and the undulation range of the second order is 0-1.②The average radius of the first experiment district of Mus norvegicus is 3.67 meters;The average radius of the strongest cluster district of Mus norvegicus is 21.00 meters;The average radius of the biggest cluster district of Mus norvegicus is 79.67 meters.The numbers of "hot spot" in different villages differ greatly.The undulation range of the first order is 2-5,and the undulation range of the second order is 0-1.③The average radius of the first experiment district of Rattus norvegicus is 3.67 meters;The average radius of the strongest cluster district of Rattus norvegicus is 14.00 meters;The average radius of the biggest cluster district of Rattus norvegicus is 68.33 meters.The numbers of "hot spot" in different villages differ slightly.The undulation range of the first order is 2-4,and there are no cluster hot spot of the second order.
(3)The "hot spot" analysis of NNH shows that there exist evident spatial clustered "hot spots" in rats in villages and clustered "disease foci" in rats of various species in villages.
8.The analysis of virus sequence shows the movement track of host animals The spatial relationship of SEO viruses carded by different host rats is described by using phylogenetic tree and MST.The results show that:
(1)The SEO viruses in different villages are relatively independent.
(2)But SEO viruses carried by host rats in conterminous areas of villages are homogeneous to some extent.
(3)The HFRS viruses in experiment districts show spatial structure to some extent.
(4)The viruses carried by host rats in the same villages show some spatial structure and spatial heterogeneity.For instance,in Dashanqian village of the first experiment district,some viruses carried by host rats form obvious spread track.
Conclusions:
1.The ecological niche of host rats in experiment districts
(1)The structure of community of host animals differ due to the differences of geographical ecological environments in different villages.But Rattus norvegicus and Mus norvegicus are still advantageous rats in villages.
(2)The time niche,the spatial niche,the food niche,and the feed niche of host rats in different micro ecological environment show different characteristics,which reveals that the ecological population structures of HFRS host animals are very complex in micro scale.
2.The distribution characteristics and related infection factors of host rats in experiment districts
In small ecological environment,the rat density changes according to season evidently,and the spatial distribution shows clustered character.Regular deratization, cement anti-rat floor and methods used by residents are the main reasons for the decrease of rat density,while breeding livestock in yards and mixing up food,clothes and other goods are the main reasons for the increase of rats.
3.Infection rates and indices of parasite acarid of different rats are different.Gamasid is the major mites in host animals in experiment district,and Rattus norvegicus is the main host of acarid.
4.The infection characteristics of HFRS virus in host rat in experiment district
In experiment district,although the rat density is low,the toxin-carrying rate is high. The toxin-carrying rate of Rattus norvegicus is higher than that of Mus norvegicus. Although the structure of host rats in villages differ greatly,the distribution characteristics of rat density in age,sex,and different season are similar.
5.The spatial epidemiology characteristics of HFRS virus in host rat in experiment district
In small site,the primary clustered range of host rats equates with the area of residents' room.The strongest cluster district centers in the area of 1-2 families;The average radius of the biggest cluster district of Mus norvegicus is 80 meters.The numbers of "hot spot" in different villages differ greatly.
6.The spatial ecology and spatial epidemiology characteristics of toxin-carrying host rats
(1)The relationship of SEO viruses in villages is obscure.
(2)SEO viruses carried by host rats in conterminous areas of villages are homogeneous to some extent.It means that rats may be infected by others during the process of migrating to open country,and bring viruses back to rooms during the way back.
(3)The HFRS viruses in experiment districts show spatial structure to some extent. That is to say the ecological barriers maybe exist between villages.
(4)The viruses carried by host rats in the same villages show some spatial structure and spatial heterogeneity.For instance,in Dashanqian village of the first experiment district,some viruses carried by host rats form obvious spread track.The results show that there are both ecological barriers and ecological relationship inside villages.
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