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Differences in Life History Traits and Quantities of DNA/RNA and Protein Polyploidy in Daphnia Methodology and effects on life history traits Kristian Alfsnes Master of Science Thesis University of Oslo Department of Biology Center for Evolutionary and Ecological Synthesis 2007 Daphnia pulex taken from “On the freshwater crustaceans occurring in the vicinity of Christiania” by Georg Ossian Sars, 1861 2 Preface The study was conducted at the University of Oslo, Norway, as a part of my master thesis. Ideas and initial proposals came from the ever-so-helpful professor Dag O. Hessen, which has been obliging in structuring and correcting the thesis, connecting me with people outside UiO and always being available for discussions. Countless hours with microscopy was done by professor Morten M. Laane, helping me with the cytogenetic analyses and lending great tips and hints for the other analyses. Great thanks to PhD-student Marcin Wojewodzic, who was working with me for the majority of this study, lending help whenever I needed, coming up with good solutions and reading through my manuscripts many times. Dziękuję! Per Færøvig patiently helped me with the flowcytometry and other laboratory work, Tom Andersen has lent great help with analyzing the data and the statistics and Jens Petter Nilssen for help with taxonomy and identification of the Daphnia. Thanks to Morten Skage at the University of Bergen for running microsatellite and mtDNA analyses and correcting my interpretation of the data, Anders Hobæk at NIVA in Bergen, for helping me with the microsatellite analysis, and compiling the phylogenetic tree for Daphnia with sequences gratefully received from John K. Colbourne at the University of Guelph. Thanks to Gerben van Geest at Nederlands Instituut voor Ecologie which accompanied me at Svalbard, helping with sampling of Daphnia and later sharing data from the lakes and ponds of Ny-Ålesund. Also thanks to the staff at the research station at Ny-Ålesund, Kings Bay and the staff and professors at UiO, especially thanks to the lunch-group at the 4th floor. Special thanks to family, friends and girlfriend which have endured me the last months, and giving me financial, moral and emotional support. Kristian Blindern, May 31st 2007 3 TABLE OF CONTENTS PREFACE ..................................................................................................................................................... 3 CONTENTS.................................................................................................................................................. 4 ABSTRACT .................................................................................................................................................. 5 INTRODUCTION........................................................................................................................................ 6 MATERIALS & METHODS .................................................................................................................... 13 1.1. SAMPLING & CULTIVATION................................................................................................................ 13 1.2. CYTOGENETIC ANALYSES ................................................................................................................... 21 1.3. LIFE HISTORY EXPERIMENT ................................................................................................................ 28 1.4. GENETIC ANALYSES ........................................................................................................................... 33 RESULTS.................................................................................................................................................... 42 2.2. CYTOGENETIC ANALYSES .................................................................................................................. 42 2.3. LIFE HISTORY EXPERIMENT ............................................................................................................... 53 2.4. GENETIC ANALYSES ........................................................................................................................... 67 DISCUSSION.............................................................................................................................................. 78 3.2. CYTOGENETIC ANALYSES .................................................................................................................. 78 3.3. LIFE HISTORY EXPERIMENT ............................................................................................................... 85 3.4. GENETIC ANALYSES ........................................................................................................................... 95 CONCLUSION......................................................................................................................................... 106 REFERENCES ......................................................................................................................................... 108 APPENDIX ............................................................................................................................................... 114 4 Abstract Four arctic, obligate parthenogenetic populations assumed to be polyploid and one temperate, cyclic parthenogenetic, diploid population were used to compare life history traits, growth rates and content of nucleic acids for polyploidy versus diploidy. Cytogenetic studies were done to evaluate the level of ploidy optically or with the aid of optical instruments. A life history experiment was run to identify variations in fitness parameters, fertility, maturation, growth and survival, identifying the adaptive effects and costs of polyploidy. Variations in nucleic acids in polyploids and diploids were accessed by quantifying DNA, RNA and protein. A microsatellite analysis was run to verify the assumed ploidy levels, and species identification was done by sequencing mtDNA and comparing with previously published sequences. The study has a strong focus on assessment of different methods and is in structure affected by this. Microsatellite analysis positively identified the arctic populations as polyploids, and the temperate as diploid. No assessment of the chromosome numbers was reached using cytogenetic analyses. Statistical difference was noted between the nuclei size in the intestine and connective tissue between the diploid and polyploid populations. Polyploid populations were found to have lower population growth, lower fertility, smaller size and delayed maturation, lower growth rate and lower survival than the diploid population. Ploidy level and quantity of nucleic levels and ratios were not found to be correlated, but RNA and DNA per dry weight and RNA/DNA ratio was found to be weakly correlated with specific growth rate when controlled for the ploidy levels. 5 Introduction Polyploidy in Daphnia has repeatedly been reported in the Arctic where it seems more wide-spread than in temperate and tropical latitudes (Adamowicz et al., 2002). It is occasionally found north of 58° N in North America and Europe, and become common at 70° N (Beaton & Hebert, 1988; Ward et al., 1994). Observations of polyploid Daphnia has also been reported in alpine locations in Argentine at 46° S (Adamowicz et al., 2002). Polyploidy in Daphnia is suggested to have arisen during the glacial-cycles in the Pleistocene; glacial expansion producing isolated refugia (selecting for parthenogenesis), glacial retreats then allowing for secondary contact and hybridization between closely allied species (Dufresne & Hebert, 1994; Dufresne & Hebert, 1997, Weider & Hobæk, 2003; see also Fig. 43). Distribution of polyploidy could thus be more based on ecological and historical contingencies than direct selection for polyploidy (Adamowicz et al., 2002). Polyploidy has been found to be correlated with increase in cell size, delay of maturity, increased size at maturation, lower fertility and increase in offspring size (Levin, 1983; Weider, 1987). Genetically, polyploidy increase the DNA-template, increasing production of RNA and protein, and the genetic variation (as each individual carries several copies of the DNA-template). However differences in life history traits are easily confounded with the concurrence of parthenogenesis breeding mode (Weider, 1987; Otto & Whitton, 2000). The advantage of increased genetic variation (heterozygosity) by increasing the genetic template available for mutations, silencing and reshuffling of duplicated genes (Bailey et al., 1978; Otto & Whitton, 2000) should be evident in obligately asexual populations, as polyploidy increases plasticity to environmental variations in populations otherwise unable to shift in response to changes (Dufresne & Hebert, 1998). Cell division is found to be impeded at low temperatures, increasing the initial genetic template would hence allow for increased metabolic activity in the early stages which otherwise would be limited by the genetic template for production of RNA and proteins (Grime & Mowforth, 1982). Polyploidy in arctic Daphnia may also be the result of hybridization events between isolated populations during the Pleistocene glaciations (Dufresne & Hebert, 1994; Dufresne & Hebert, 1997). 6 The study aims to determine the ploidy level in temperate and arctic populations. Comparing life history traits and nucleic contents of these populations to examine the effect of increased ploidy levels and draw some conclusions of possible causations of the observed increase in polyploidy
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