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Ebert D. Ecology, Epidemiology, and Evolution of Parasitism in Daphnia [Internet]. Bethesda (MD): National C enter for Biotechnology Information (U S ); 20 0 5 .
Bookshelf ID : NBK 2 0 3 9

Acknowledgments
In 19 9 0 , I w as w ork ing on my PhD on the life- history evolution of Daphnia at the Z oological Institute of the U niversity of Basel w hen Bill H amilton came for a visit there. Being deeply interested in the evolution of parasites, he did not w ait long before ask ing me about parasites in Daphnia. T o my embarrassment, I had to answ er that I had never seen a parasite in Daphnia and that I believed they may not be very important. O f course Bill w as not convinced by my obviously rather superficial and uninformed statement, and after some discussion I agreed that I w ould collect some field samples the nex t day and screen them w ith him for parasites. Ex cept for a few individual Daphnia carrying some algae epibionts, w e did not find anything. Bill w as convinced, how ever, that there w as more to discover, and a month later he sent me a copy of G reen' s 19 7 4 paper on " Parasites and Epibionts of C ladocera." G reen lists numerous symbionts living on and in Daphnia and other cladocerans too many to be ignored. A t the same time, Paul S chmid- H empel s group w as w ork ing at the Z oological Institute in Basel, establishing bumblebees as a new model system to study hostparasite interactions. O n the basis of many interactions w ith Paul and his students, mainly J acq ui S hyk off and C hristine M ller, I started to develop an interest in host parasite evolution and eventually decided that this may be a good topic for a postdoc. Interestingly, other PhD students in our group pursued the same direction. Being an ex perimentalist at heart, I decided that I w ould lik e to start w ork ing w ith a rather simple hostparasite system that w ould allow me to do sophisticated ex periments. I ask ed Bill H amilton to host me in O x ford. Bill w as enthusiastic, but w e w ere undecided about w hat system w ould be good. Bill suggested Daphnia, but I w as not so sure, because I still had not seen parasites in Daphnia. T he advantage w ith Daphnia, how ever, w as that I already had ex perience w ith them and that they seemed, indeed, lik e a good system: easy to maintain, reproducing clonally, short generation time, and small siz e, to name j ust a few . Nonetheless, before I settled on Daphnia, I did an intensive literature search (back in the early nineties, this w as still a time- consuming enterprise) and came up w ith three host systems I w anted to check out: the Indian meal moth (P l o d ia int e r pu nc t e l l a), T r ib o l iu m , and Daphnia. T he tw o earlier systems are genetically more tractable than Daphnia, because genetic crosses of Daphnia clones are still not very easy. In the end, one factor sw ayed my decision to w ork w ith Daphnia: H ow could one control airborne pathogens during ex periments in w hich controls and infected replicates are standing side by side? W ith w aterborne pathogens, I reasoned, this w ould be much less of a problem. It turned out to be correct. My initial field surveys around O x ford w ere not very successful: plenty of Daphnia, but no parasites. A fter several months, I decided to contact J im G reen, w ho w as still w ork ing in L ondon but w as about to retire. H e invited me to bring him the samples in w hich I w anted to find parasites. It took him 10 minutes to find the first parasites, and over the nex t tw o hours a few more species turned up as w ell. W hat I did not k now w as that it is essential to use phase- contrast microscopy, because microsporidian parasites, in particular, are barely visible w ithout it. Despite his enthusiasm, J im w as rather pessimistic about the possibilities of culturing Daphnia parasites and conducting controlled ex periments. H ow ever, back in O x ford I check ed all of my cultures and realiz ed that I had already cultured one species for several months together w ith the D. m ag na clone it lived in. W ithin 2 w eek s I w ork ed out a method for controlled culturing of G l u g o id e s int e s t inal is (formerly P l e is t o pho r a int e s t inal is ) and 3 months later submitted the first paper on it. S ince then, my lab has developed culturing methods for more than 10 species of parasites. T his w ork w ith the Daphnia parasites has absorbed my attention since then, and together w ith my research group, I have spent many ex citing hours in the laboratory and in the field finding out more about Daphnia parasites, and about parasites in general. T he Daphniamicroparasite system has proved to be a pow erful model for many q uestions in basic epidemiology, ecology, and evolution. T he upcoming genome data w ill ex tend this into the Narrow R oads of G ene L and (the title of Bill H amilton s second book w ith many parts on the evolution of hostparasite interactions, 20 0 1) and w ill hopefully open up a pluralistic approach to understanding hostparasite coevolution. T his book w ould not have been possible w ithout the collaboration of the marvelous people w ork ing w ith me in O x ford, S ilw ood, Basel, and F ribourg, and also collaborators from around the w orld w ho shared my enthusiasm for hostparasite interactions. I w ant to thank the people in and around my group (in roughly chronological order): V alentino L ee, K atrina Mangin, H eide S tirnadel, S ven K rack ow , J udy W earing- W ilde (O x ford), Dermot McK ee (S ilw ood Park ), C hristine Z schok k e- R ohringer, H ans- J ochaim C arius, T om L ittle, Daniel F els, Marc C apaul, Myriam R iek , Patrick Muck low , Pia S alath , K atj a Pulk k inen (Basel), C hristoph H aag, Dominik R efardt, Dita V iz oso, O lga S ak w insk a (Basel and F ribourg), L usia S ygnarsk i, S andra L ass, Marc Z binden, K nut H elge J ensen, R affael A ye, F lorian A ltermatt, H olly G anz (F ribourg and Basel), and T homas Z umbrunn (Basel). S pecial thank s to J rgen H ottinger, w ho became over the years not only a close friend but also the irreplaceable center of the group. Numerous collaborators, many of w hom are theoreticians, helped open my eyes w hen I w as blinded by the beauty of Daphnia symbionts: S ebastian Bonhoeffer, Marc L ipsitch, Martin Now ak , W olfgang W eisser,

R ichard L ensk i, Bill H amilton, Paul R ainey, Mitj a S cholz , Martin Embley, J anne Bengtsson, L iz C anning, S teve S tearns, Ilmari Paj unen, R oland R egoes, R onny L arsson, K erstin Bittner, Ellen Decaesteck er, and Paul S chmidH empel. F lorian A ltermatt, H olly G anz , S andra L ass, Dominik R efardt, Marc Z binden and T homas Z umbrunn read most of the book in earlier drafts and helped me to polish style and content. Dita V iz oso produced several figures to illustrate important aspects of life cycles and transmission. Dita V iz oso and F rida Ben- A mi are thank ed for contributing photographs. I thank my son G leb for help w ith producing some of the figures. S uz anne Z w eiz ig improved the style and readability of the language throughout the book . R ita G unasek era helped me w ith the formatting of the various files. J o McEntyre and L aura Dean w ere of great help in all aspects of publishing this book . T hank s to their w ork , the book can be made available to a w ide audience via the Internet. F inally, I w ant to thank my friends and family for their support and the encouragement to w rite this book .

References

1. G reen J , 19 7 4 . Parasites and epibionts of C ladocera. T ransactions of the Z oological S ociety of L ondon; 3 2:4 17 - 5 15 . 2. H amilton W D, 20 0 1. Narrow roads to gene land. V ol. 2, Evolution of sex . O x ford: O x ford U niversity Press. 20 0 5 , Dieter Ebert.

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