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Lyme Disease and Biodiversity Caren Collins, Lily Hoffman-Andrews, Amanda McLoughlin, and John Soghigian Advisor: Dr. Todd Livdahl Modeling the System The Infective Agent and its Vector: In order to test the host dilution concept, a model was constructed using the software, STELLA. The model contained parameters for nine species and Ixodes scapularis. (fig 1). Figure 2 is a close up of the interactions between tick and whitefooted mouse. The model was run with different community compositions, varying from a low biodiversity community to higher biodiversity community, with nine species. Figure 3 shows a community, in which the tick has only the white-footed mouse as prey. Figure 4 demonstrates a community that is composed of the ticks, mice, and squirrels. Figure 5 shows how a very low density of mice infected with Lyme disease can rapidly spread the disease. Figure 6 shows a community with nine species, a representation of host dilution. Borrelia burgdorferi is the spirochete bacteria responsible for Lyme Disease (Fig. 1). It resembles other spirochetes in that it is a highly specialized, motile, two-membrane, spiral-shaped bacteria which lives primarily as an extracellular pathogen. In 1982, Willy Burgdorfer isolated Borrelia burgdorferi from Ixodes ticks, as well as from patients with Lyme disease. Ticks are small arachnids that act as ectoparasites, living on the blood of mammals, birds, and even reptiles (Fig. 2). Ixodes scapularis is the vector in the Northeast and North central United States. Ixodes pacificus is the vector in the Northwest United States. Species Body Burden White-footed mouse The tick life cycle requires two years and has three feedings with three stages. Adult ticks mate in the fall and early spring and the females lay their eggs on the ground. These eggs hatch into larvae by summer. The larvae then feed on small animals, particularly the white-footed mouse, in the summer and early fall (first feeding). The larvae are inactive until spring when they molt into nymphs. Nymphs feed on small animals in the late spring and summer (second feeding) and molt into adults in the fall. The adults then feed, usually on white-tailed deer, and the cycle begins again. Ticks become infected with Borrelia burgdorferi when they feed on infected animals. Ticks can become infected at any of the three feedings and can then pass on the bacteria to animals and humans during subsequent feedings. C TF Sor ex 1390 20.61% 36 55 25 900 13.35% White-tailed deer 239 4.6 0.25 59.75 0.89% Raccoon 127 1.3 0.2 25.4 0.38% Virginia opossum 254 2.6 1 254 3.77% Stripped Skunk 66.8 9.7 0.05 3.34 0.05% Short-tailed shrew 62.9 41.8 25 1572.5 23.32% Sorex Shrew 55.5 51.2 25 1387.5 20.58% 142 14.7 8.1 1150.2 17.06% U ninfected D ee r U ninfected Sor ex U ninfected D ee r D ea ths So rex D R In fected Dee r D ea ths U ninfected Opo ss um N/A A map was made of small mammal biodiversity in the continental United States (Figure 10), using Idrisi’s Land Change Modeler, to determine whether there is a relationship at this scale between biodiversity and Lyme disease. Figure 11 and Figure 12 show average species richness and 2005 Lyme disease rates, by state. In fected Opo ss ums U ninfected Opo ss um DR Opos su mD R In fected Sor ex DR Figure 10 – Small mammal species richness in the United States. Opos su mIn fec tion s Opos su mBirth s U ninfected Sor ex DR N/A Opos su mBody Bu rde n In fected Sor ex So rex In fection s So rex Bi rths D eer Infe ctio ns ~70,000 Opos su mC ompe tenc y R ate Opos su mBR In fected Dee r D eer Bir ths C TF Opo ss um N/A So rex Bo dy Bu rde n So rex BR D eer Bir th R ate N/A So rex C omp ete nc y R ate D eer Co mp ete ncy Ra te % Chance for Tick 50 Ixodes C TF D ee r Ticks/Hec/Species 92.1 Red & Grey Squirrel Fig. 2 Ixodes scapularis Density Per Hectare 27.8 Eastern chipmunk Fig. 1 Borrelia burgdorferi Reservoir Competence GIS – Biodiversity and Lyme disease rates In fected Opo ss um DR D eer De ath Ra te D eer Bo dy Bur den tic k b irth rate C TF ST Shre w C TFSqu irre l In fected Tick s Sq uir rel Compe ten cy Ra te U ninfected Tick s ST Sh rew C ompetenc y R ate Sq uir rel Birth R ate ST Sh rew Bo dy Bu rde n ST Sh rew BR tic k b irth s U ninfected Sho rt Tail Shr ew In fection U ninfected Squ irre l More De ath s D eath s Sq uir rel Infe ctio ns Sq uir rel Birth s U ninfected Sho rt Ta il Shre w Birth s In fected Squ irre ls In fected ST Shr ew D eath R ate ST Sh rew In fec tion s Sq uir rel Bod y Burd en U ninfected Squ irre l D eaths In fected Squ irre l D eaths U ninfected ST Shr ew De aths In fected ST Shr ew DR Sq uir rel Dea th Ra te Sh ort Ta il Sh rew D R Grap h 1 C han ce to fi nd Mic e Mous e C ompe ten cy R ate C TF R ac co on R acc oo n C ompetenc y R ate C TF C hi pmu nk Mice Birth R ate C hipmun k C ompe ten cy Rate C TF Sku nks Sk un k C ompetenc y R ate C hipmun k BR Sk un k Body Bu rde n Stripp ed Sku nk BR U ninfected Mic e R acc oo n Bi rth Ra te In fected Mic e U ninfected C hi pmunk s In fected Chi pmunk s U ninfected Strip pe d Skun k U ninfected R ac co on In fected Rac co on s Mice Infe ctio ns Mice Births C hipmun k Birth s Stripp ed Sku nk Bir ths R acc oo n Bi rths In fected Sku nk C hipmun k In fec tio ns Stripp ed Sku nk Infecti ons R acc oo n In fec tion s Mice De atbs Mous e Body Bu rd en U ninfected R ac co on De aths In fected Rac co n D ea ths Mous e D ea th R ate Mice De aths U ninfected C hi pmunk De ath s C hipmun k D ea th R ate In fected Chi pmunk De ath s U ninfected Strip pe d Skun k D ea ths Stripp ed Sku nk DR In fected Sku nk DR C hipmun k Bod y Burd en R acc oo n D eath R ate R acc oo n Bo dy Bu rde n Figure 11 Figure 12 Fig. 4: STELLA Model Fig. 5: Close up of the model: the tick sector and a species sector Figure 12 Fig. 3 Tick Life Cycle A statistically significant negative correlation between average biodiversity and Lyme disease rates was found (Pearson’s r = -0.338, N = 48, p = 0.019). The results of the non-linear regression are shown in Figure 12 and Table 1. While these results must be taken with some caveats, given the very broad scale of the analysis, they provide intriguing support for the host dilution hypothesis, especially when taken along with our model results. Table 1 Host Dilution The Host Dilution hypothesis predicts that in communities with high species diversity, the effect of the primary reservoir of a disease, such as the white-footed mouse in Lyme disease, can be diluted by the presence of other, less competent hosts of the disease. These other hosts allow for the feeding of the vector, in this case, the tick, but rarely infecting the vector or becoming infected. In the Host Dilution hypothesis, there are different roles that the each organism in the community has. We define dilution hosts as those characterized by relatively high body burdens with low reservoir competence and a high population density. In the case of Lyme disease, species such as squirrels act as dilution hosts in the population. Rescue hosts are those that have relatively high body burdens with moderately high reservoir competency and are capable of maintaining the disease in the population when the most competent host density is low. Shrews would be an example of a rescue host in this community. Fig. 6: Simulation run with infected ticks and initially uninfected mice Fig. 8: Simulation run with infected mice and initially uninfected ticks. Fig. 7: Simulation run with squirrels and mice References Animal Diversity Web. Accessed December 18, 2008 at http://animaldiversity.ummz.umich.edu. Clark Labs. 2006. Idrisi Land Change Modeler. Computer software. Clark Labs, Worcester, MA, USA. LoGiudice K, Ostfeld RS, Schmidt KA, Keesing F. The ecology of infectious disease: Effects of host diversity and community composition on Lyme disease risk. Proceedings of the National Academy of Sciences 100(2): 567571. Ostfeld RS, Keesing F. 2000. Biodiversity and disease risk: The case of Lyme disease. Conservation Biology 14(3): 722728. Patterson, BD, Ceballos G, Sechrest W, Tognelli MF, Brooks T, Luna L, Ortega P, Salazar I, Young BE. 2007. Digital Distribution Maps of the Mammals of the Western Hemisphere, version 3.0. NatureServe, Arlington, Virginia, USA. Accessed at http://www.natureserve.org/getData/mammalMaps.jsp on 12/1/2008. Penn State College of Agricultural Sciences. Lyme Disease: Life Cycle of Blacklegged Tick. 11 September 2001. http://www.ento.psu.edu/Lyme/lifecycle.htm> Science Daily. Ticks Don’t Come Out in the Wash. 11 October 2007. http://www.sciencedaily.com/releases/2007/10/071006083356.htm Wadsworth Center, New York State Department of Health. Disease Carriers, Bacteria: Borrelia burgdorferi. http://www.wadsworth.org/databank/borreli.htm Fig. 9: Simulation run with all nine species. An example of biodiversity at work.