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Genetic Variation and Inbreeding Depression in the Rare California Endemic, Astragalus Agnicidus (Leguminosae)

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Genetic Variation and Inbreeding Depression in the Rare California Endemic, Astragalus Agnicidus (Leguminosae) Genetic Variation and Inbreeding Depression in the Rare California Endemic, Astragalus agnicidus (Leguminosae) by Robin Bencie A Thesis Presented to the Faculty of Humboldt State University In Partial Fulfillment of the Requirements for the Degree Master of Arts December, 1997 GENETIC VARIATION AND INBREEDING DEPRESSION IN THE RARE CALIFORNIA ENDEMIC, ASTRAGALUS AGNICIDUS (LEGUMINOSAE) by Robin Bencie We certify that we have read this study and that it conforms to acceptable standards of scholarly presentation and is fully acceptable, in scope and quality, as a thesis for the degree of Master of Arts. Michael R. Mesler, Major Professor Mich J. Bowes, Commit tee ember Timothy E. Lawlor, Committee Member Andrea J. Pickart, Committee Member John O Sawyer, Committee Member Milton J. Boyd, Graduate Coordinator Linda A. Parker Dean for Research and Graduate Studies ABSTRACT Predictions of low genetic variation and low levels of inbreeding depression in small plant populations were tested on Astragalus agnicidus, a species with only one known population. Loss of alleles through drift and increased inbreeding (followed by selection) may leave small populations genetically depauperate, but with negligible genetic load as well. This scenario results in low levels of inbreeding depression as selfed and outcrossed progeny become more equivalent in overall fitness (i.e., the difference in progeny fitness between pollination treatments decreases). As predicted, gel electrophoresis of isozymes indicated that A. agnicidus has low genetic variation. Five fitness variables: seed set, seed weight, germination, survival, and seedling weight were used to test for fitness differences between open-pollinated and self-pollinated progeny. Some degree of fitness reduction for selfed progeny was seen in all variables, but only in seedling survival was the difference significant. Results indicated that the effect of the maternal parent was more important than pollination treatment on determining the fitness of an individual. Therefore, there is low enzyme diversity within the population, but genetic load is still maintained in the population and varies between individuals. The role of the mating system in influencing the level of genetic variation and inbreeding was also examined. One polymorphic enzyme, IDH, was used as the genetic marker in estimating an outcrossing rate of .59, which indicates a mixed mating system. The advantages of a mixed mating system include maintenance of moderate levels of heterozygosity (and thus, genetic variation) through outcrossing and elimination of genetic load and retention of optimum genotypes through selfing. Evolutionary models predict that A. agnicidus, with a high level of self-compatability and an inbreeding depression level of <50% would evolve towards a predominantly selfing mating system. However, A. agnicidus's crossing rate remains intermediate due to an obligatory, insect mediated, tripping mechanism that virtually assures some cross- pollination in the system. iii AKNOWLEDGMENTS I am grateful to those who supported me during the course of this project. Andrea J. Pickart, Area Ecologist for The Nature Conservancy, was instrumental in literally bringing this project, and thus graduate school, into my world. She secured significant grant funds from The Hardman Foundation and The Department of Fish and Game's Endangered Plant Program. We visited over statistics and the history of Astragalus agnicidus. My major professor, Michael Mesler, has always been my botany mentor, and his tremendous support and encouragement throughout this project is appreciated deeply. My fellow graduate students were =hesitantly willing to listen, and always understood (even Gordon, most of the time). The Biology Department's staff and faculty were very generous with their time and expertise. The Tosten family graciously gave me unlimited access to the site of A. agnicidus. And, most of all, I give thanks to plants. iv TABLE OF CONTENTS Page No. I. ABSTRACT iii II. ACKNOWLEDGMENTS iv III. LIST OF TABLES vi IV. LIST OF FIGURES vii V. INTRODUCTION 1 VI. THE SPECIES 4 VII. OBJECTIVES 9 VIII. METHODS 11 Genetic variation 11 Mating system 11 Inbreeding depression 13 IX. RESULTS 16 Genetic variation 16 Mating system 19 Inbreeding depression 22 X. DISCUSSION 28 XI. CONCLUSION 33 XII. RECOMMENDATIONS 34 XIII. LITERATURE CITED 36 LIST OF TABLES Page No. Table 1. Summary of gel electrophoresis data 18 Table 2. Allele frequencies, inbreeding coefficients, and results of tests for random mating 22 Table 3. Average values for progeny fitness variables ................... 24 Table 4. Results of pollination treatment and maternal parent on progeny fitness variables 26 Table 5. Relative performance (RP) values for maternal parents 27 Table 6. Genetic diversity comparisons between Astragalus agnicidus and other species with similar life history traits 30 vi LIST OF FIGURES Page No. Figure 1. Vicinity map of the Astragalus agnicidus site 5 Figure 2. Map of subpopulation locations 6 Figure 3. Electrophoretic banding patterns for IDH 18 Figure 4. Electrophoretic banding patterns for 6-PGD 20 vii INTRODUCTION The only known population of the Humboldt milk-vetch, Astragalus agnicidus Barneby, provides one example of the possible genetic and fitness scenarios that can occur in plant populations with few individuals. Small, isolated populations are vulnerable to severe reductions in fitness due to inbreeding and loss of genetic variability (Barrett and Kohn, 1991; Ellstrand and Elam, 1993). Rare plants that occur as fragmented populations can be especially vulnerable to extinction, as any single population's success may be pivotal in preservation of the species. Yet in spite of the potential negative consequences of small population size, some rare plants maintain relatively high levels of genetic variability or exhibit low levels of inbreeding depression (Karron, 1987; Ellstrand and Elam, 1993; Richter, 1994). This study investigates the degree of genetic variability and the amount of inbreeding depression present in a rare, California endemic plant. The role of the mating system in maintaining genetic variability is also examined. My intention is to use this information to formulate conservation guidelines that will sustain, and potentially improve, the health of the population. This study also provides empirical evidence for testing general predictions made by theoretical models about the relationship between inbreeding depression, the selfing rate, and the evolution of plant mating systems (Lande and Schemske, 1985; Holsinger, 1988). After suspicions of it's toxicity to livestock provoked the systematic removal of all individuals from the site, A. agnicidus was presumed extinct (Berg and Bittman, 1988). Although the species later recovered from a dormant seed bank, the progressive decrease in the existing population size has made A. agnicidus highly susceptible to loss of genetic variability via genetic drift and mating between closely related individuals (biparental inbreeding). Genetic variation is considered crucial for long-term survival as it provides the necessary genetic foundation for adaptation to changing environmental conditions (Grant, 1991). Although some rare plants benefit from reduced genetic variation in the form of highly specialized genotypes, these relatively short-term benefits 1 2 may limit a population's ecological range and long-term genetic flexibility (Barrett and Kohn, 1991; Huenneke, 1991). Genetic drift can cause dramatic fluctuations in allele frequencies in small populations, leading to allele fixation in just a few generations. The rare and infrequent alleles that enhance genetic diversity, and that may ultimately code for traits that are necessary for survival in response to environmental change or crisis (i.e., disease or predation), face the greatest probability of extinction by virtue of their low frequency. While a single, undivided population can quickly lose alleles, A. agnicidus 's isolated subpopulations may offer some protection against loss of allele diversity as subpopulations randomly diverge (Hartl, 1987; Barrett and Kohn, 1991; Grant, 1991; Ellstrand and Elam, 1993). In small populations, inbreeding can rapidly lead to the expression of lethal and deleterious alleles in homozygous genotypes, and a concomitant reduction in fitness and allele diversity. If a population can survive this initial inbreeding depression that comes with purging detrimental alleles (genetic load), then the negative consequences of selfing are significantly diminished. Assuming A. agnicidus is a self-compatible species that has existed historically as a small population, then a low level of genetic variability, genetic load and inbreeding depression would be expected. However, if this population has only recently experienced a substantial reduction in size (bottleneck), then the level of genetic load and potential for inbreeding depression may still be high and pose a significant threat to the above ground population (Barrett and Kohn, 1991). The amount of inbreeding depression has been considered one of the primary factors influencing the evolution of plant mating systems. Genetic models have shown that when the fitness level of selfed progeny is < 50% of the fitness level of outcrossed progeny (high inbreeding depression), a predominantly outcrossing mating system will be maintained (Fisher, 1941; Lande & Schemske, 1985). Alternatively, if the fitness level of selfed and outcrossed progeny is more
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