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A Genetical Study of Isolated Populations of the Australian Bush

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A Genetical Study of Isolated Populations of the Australian Bush 4t"11 A GENETICAI SI1]DY OF ISOLATED POPULATIONS OF ]IIE AUSTRALIA}I BUSH-RAT, RATIUS FUSCIpES by Líncoln II. Sclunitt, B. Sc . (Ilons . ) DeparËment of Genetics Universlty of Adel-aide A thesis subrnitted to the Universíty of Adelaide for the Degree of Doctor of Phllosophy in October, L977. A,,a-cüd Ì1"¡ cA lq /'i TASLE OF CON'IENTS Page SI]MMARY J- DECI,ARATION r_aa ACKNOWI,EDGEMENTS 1v CIIA?TER 1 INTRODUCTION I CHAPTER 2 GENETIC VARIATION IN NATURAI POPULATIONS 5 2.1 Ge1 electrophoresís and the frequency of polynor phisms 5 2.L.1 Límitatíons of the techníque 6 2.L.2 Levels of varíabilíty in different taxa 7 2.r.3 Levels of variabílity in different proteins 8 2.2 Geographic variation ín gene frequencies 9 2.3 Forces responsible for genetic polymorphisms 10 2.3.1 Selectíve replacement of a gene 10 2.3.2 Genetic dríft 11 2.3 .3 HeterozygoËe advantage 11 2.3.4 Frequency dependent sel-ection 13 2.3 .5 Het erogeneous envír onments t4 2.3.6 Natural selection Ðersus the neutral- mulation-random drift theory 15 2 .4 Genetic variation in island populaÈions 16 2.4.I The fevel of heterozYgositY 16 2,4.2 Other aspeets of island populatíons L9 CHAPTER 3 TTIE BIOLOGY OF RATTUS ÍUSCIPES 23 EIE CTROPHCRETI CALLY DETE CTABLE VARTATTON CHAPTER 4 MATERIAIS AND EXPERIMENTAL METTIODS 28 4.1 CollecÈíng localitíes and techníque 28 4.2 MaÍntenance of laboratory colonY 28 4.3 Tissue extraction procedures 29 4.4 ElecErophoretíc procedures 30 4.4.t Acid phosphaÈase 32 4.4 .2 Albumín 33 4.4.3 Esterase 33 4 .4.4 Glucos e-6-phosphatç d ehydro genas e 34 4.4.5 Glutamate oxaloacetat e transamínase 34 4.4.6 Haemoglobin 34 4.4.7 Is ocítrate dehydrogenase 35 4.4.8 Lactate dehydrogenase 35 4.4.9 lIalate dehydrogenase 35 Page 4.4.10 Malfc. enzyme 36 4.4.1I Phosphoglucomutase 36 4. 4.12 6-Phosphogluconate dehydrogenase 36 4.4.I3 Superoxide dismutase 36 4.s Subcellular localisatíon 37 4,6 Nomenclature 37 CIIAPTER 5 RESULTS 39 5.1 Phenotypes, famlly data and geographíc varíatíon for each proËeín 39 5.1.1 Acid phosphatase 39 5,1.2 Albunín 39 5.1 .3 Esterase 40 s.1.4 GlutamaLe oxaloaceËate transamínase 4l 5.1.4.L Supernatant glutamate oxaloacetate transaminase 4L 5. 1.4.2 l"Iitochondríal glutamate oxaloacetaËe trans aminase 43 5.1 .5 Glucose -6-phosphate d ehydro genas e 44 5.1.6 Haemoglobín 44 5.1.7 IsocitraËe dehydrogenase 47 5.1.7.1 Supernatant isocitrate dehydrogenase 47 5.1.7 .2 Mitochondrial isocítrate dehydrogenase 48 5.1.8 Lactat.e dehydro genase 49 5.1.8.1 l-actate dehydrogenase: B subunit 51 5. 1 .8 .2 Lactate dehydrogenase: A subuníc 51 5.1 .9 Malate dehydrogenase 52 5.1 .10 Malic enz)rme 53 5.1 .11 Phosphoglucomrtase 55 5.I.12 6-Phosphogluconate dehydrogenase 57 5.1.13 Superoxide disumtase 5B 5.2 Suunnary of geographic variation 58 5.3 Estimates of varíaEion within populations 60 5.4 Intrapopulation ass ociations 61 5.5 MulËívaríate analyses of isozymic data 62 5.5.1 Genetic dístance 62 s.5.2 Correspondence analysis 66 5.5.3 Genetic distance, geograPhic distance and the tíme since isolation 67 CHAPTER 6 DISCUSSION 7L 6.L Soluble glutamaËe oxaloacetate transamínase in the Pearson Islands 7T 6.2 Geographic varíation in mítochondrial isocítrate dehydrogenase 73 6.3 Genetic variabílity hriËhin populations 74 6,4 Geographlc variation in gene frequencies BO 6.5 l"lultivariate analyses of genetic data 82 6.6 Genetic dísËance, taxonomíc rank, and Èhe' relat ionship between subsPecies 87 6.7 Evolutionary forces 90 6.8 FuËure sÈudies 93 Page METRICAT, VARTATTON CHA?TER 7 METR IC VARIATION IN RATTUS FUSCTPES GREYTÏ 96 7.L InËroductíon 96 7.2 Materials and experjmental methods 97 7.3 StatisËical methods 100 7.4 Results 101 7.5 Discussion 103 APPENDIX 1 NO]ES ON TIIE ECOLOGY AND REPRODUCTION OF RATTUS FUSCIPES GNUvil APPENDIX 2 PUBLISHED PAPERS 1. GeneËic evídence for the existence of two separate populations of Rattt'ts fuseipes gteyíí on Pearson Island, South Australia. 2. Genetic varíatíon in isolated populations of the ArrsËralían bush-rat, Rattus fuscípes. 3. l"litochondr:ial íso-citrate dehydrogenase variation in the Australían bush-rat, Rattus fuscipes greyii. 4. An elecÈrophoretíc invesËígaËion of the bindíng of 3-rkc coumarin to rat serum proteins. ABBRNVIATIONS BIBLIOGRAPTTY l_ SUMMARY A study has been made of electrophoretícally detectable protein variation in fourteen isolated populatíons of the Australian bush-rat, Rattus fusei'pes çr'eyii. A small number of índividuals from each of the other three .R. fuscipes subspecies Í/ere examined. Farníly daËa f rom abouË 50 laboratory matíngs suggest that the thirteen proteins examíned represent the products of genes aÈ síxteen different loci The Æ. f. gneyü populations inhabít several small off-shore islands (less than 300 ha), a large ís1and (about 400,000 ha), and three separate mainland areas (each about 400r000 ha). Seven sma11-island populations are monomorphic for all loci and two other sma11-island populatíons are each polynorphic at one l-ocus on1-y. The mean heterozygosity for Èhe snall-island populaËions is 0.007. These populations are genetÍcally less varíable than most manmalian populatíons. The three mainland and the large-ísland populatíons are polyrnorphíc at two to four genetic loci and the mê,an heterozygosíty in these populations is O.O42. .The .R. f, gveAíí populations studíed are reproductively isolaÈed from one another and the tíme of ísolation can be esËimated at between 5,000 and 141000 years. It ís presumed that prior to ísolation these populations shared a cormnon gene pool. The variaËion at several genetic locl throws líght on Èhe composítion of this ancestral gene pool-. Some genes, found only in one population, are apparently recent muËanÈs, while others are more widespread, indicaÈing Ëhat they were probably present in the ancestral population. For some loci ít seems líkeJ-y that cl-ines in gene frequency \¡rere present in the ancestral populaÈion. These inferences are supported by evidence from protein variaËion in the three other R. fuseipes subspecies. It is suggested that in the past,.R. f. greAií has been a link between the eastern and r'restern Australían subspecies of R. fuscipes 1i Ttre gene frequency data are sr:mmarised by a modifícation of the method of principal componenÈ analysis for contingency tables and by other genetíc distance measures. The genetíc disÈances between R. f. greA¿i populations are on average, greater Ëhan Èhe genetÍc dístances reported hítherto between oËher maumalían populatíons belongíng to the same subspecies and the genetic dístances between R. fuseipes subspecies are greater than the distances between other manmalían subspecíes. The genetic dissÍmi1-aríty beÈween -R. f . greyii populatÍons appears to have evolved exËremely rapídly, by the fixatÍon of different genes present in the ancestral populaËion and by the incorporation of ne\¡I mutants since isolation. There is evídence thaË both natural selectíon and random genetic drift have played important roles in determÍning Ëhe gene frequencies in the R. fuseLpes populations. A study of metrícal variaÉion of eighteen skul1 and body characters ín R. f. greAii populatíons revealed signif icant heterogeneíty betr^reen populations for all characters. For most characters there is no signífícant difference between.small ísland and large popul-aËions. The variance ín each character is less ín small populations than ín large populatíons, but this dífference Ín most cases is not statistically significant. The relationshíps between the populations as described by multívaríate analyses of the metric daËa is siurí1ar to that described by the proteín variation. ffi DECLARATION I declare that this thesls contains no materfal_ whÍch has been accepted for the award of any other degree or díplonra in any University, and that to the best of my knowledge and belief, íË contaíns no unteríal prevlously published or writ,ten by another person, Ð(cepÈ where due reference is made. L. H. Schmítt fv ACKNOÌ.ILEDGM{ENTS I am sincerel-y grateful to Dy supervisors, Professor J.H. Bennett, Dr. D.L. Hayrnan and Dr. R.lf. Hope, for theír helpful suggestíons, constructive criticisms and assistance r^7íth this work. In acldiËíon, Mr. J.G. oakeshoÈt and Mr. R.B. Hal-liday provided much help and discussion. Dr. R.J. trrlhite kindly provided statistical advi-ce and computed the canonícal discríminant and correspondence analyses. Dr. G.C. Kirby, Dr. P.R. Baverstock, Dr. C.R. Thridale and Dr. M. Bul1 also rnade valuable comments. Dr. N. tr{ace provided unpublished sea-bed contour maps for the area around Dog and Goat rsl-ands. Mr. P.F. Aitken and lulr. J. Seebeck provid.ed information on the dj-sÈríbutiol of -R. f. greyi¿. Dr. D. Day helped me develop the cell fractionation techniques. Technical assÍstance and advice v/as generously given by Mr. r.R. Goodwins, ÙIr. J.R.B. l,Ialpole, ft. c. chesson, Mrs. D. Goldíng and Mr. A. Kawenko. I would líke to thank Mi-ss G. Psaltis for her excellent typíng. I am indebted to Mr. J. Forrest for col-lecting the llopkíns Island sample and Dr. A.C. Robinson for colIecËing all specÍrnens other than R. f. greyií. In addition, valuable contributíons were made by the followíng people and groups who helped in many dífferenË ways while r was collecting specimens: Mr. P.F. Aitken, the Australían Department of Transport, l"lr. R. Baker, Mr. H. Enge, the Fíeld Naturalists Socíety of South Australia Mammal Club, Mr.
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