模里西斯果蠅與擬黃果蠅間隱性雌選擇的數量性狀基因分析

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研究生：	李政霖 Cheng-Lin Li
論文名稱：	模里西斯果蠅與擬黃果蠅間隱性雌選擇的數量性狀基因分析 Quantitative trait loci analysis of cryptic female choice between Drosophila mauritiana and D. simulans
指導教授：	丁照棣 Chau-Ti Ting
口試委員:
學位類別：	碩士 Master
系所名稱：	生命科學暨醫學院 - 分子與細胞生物研究所 Institute of Molecular and Cellular Biology
論文出版年：	2006
畢業學年度：	94
語文別：	英文
論文頁數：	69
中文關鍵詞：	隱性雌選擇、模里西斯果蠅、擬黃果蠅
外文關鍵詞：	cryptic female choice, mauritiana, simulans, QTL
相關次數：	點閱：2 下載：0
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同種精子優佔權(conspecific sperm precedence)是隱性雌選擇(cryptic female choice)和精子競爭(sperm competition)共同作用的結果。在生殖前隔離不完全的物種之間，同種精子優佔權可以進一步確保物種不會彼此雜混。在先前的實驗設計當中，大多數都著重在雄果蠅的遺傳變異如何影響精子競爭的結果，然而精子競爭實際上發生在雌果蠅生殖管道之中，隱性雌選擇(cryptic female choice)早已預先決定好不同雄果蠅的精子之間優勝劣敗的關係。試問隱性雌選擇是用什麼機制決定精子競爭的規則?所參與的基因又有哪些?為了能進一步瞭解其遺傳基礎，我對於D. simulans和D. mauritiana進行數量性狀基因座的定位分析，結果發現了兩個顯著影響BC1 (backcross generation one)雌果蠅生殖力(fecundity)的數量性狀基因座在X染色體上，其中可能含有雜交雌果蠅不孕的基因，另外也發現一個顯著和一個邊緣顯著影響隱性雌選擇的數量性狀基因座在第三條染色體上，和隱性雌選擇有關的基因可能就座落在這兩個基因座附近，其中嗅覺受器(olfactory receptor)是可能的基因，嗅覺受器若表現在精子、卵子或雌果蠅生殖管道可以用來對化學分子或荷爾蒙產生反應。進一步比較先前的遺傳研究，可以發現和雄果蠅性徵(male traits)或雌果蠅性選汰(female mate choice)有關的數量性狀基因座彼此相互重疊，這兩者之間的遺傳機制有連鎖不平衡(Linkage disequilibrium)的現象以及雌果蠅性選汰(female mate choice)受到後交配(postcopulatory)機制所左右都是可以進一步在第三條染色體上所發現的數量性狀基因座上加以研究的課題。

Cryptic female choice and sperm competition cooperate with each other could result in conspecific sperm precedence (CSP). While the premating isolation between two sympatric populations is not complete, CPS is important to maintain species as distinct entities. Most of previous experimental designs are focus on genetic variations among males contributing to the consequence of sperm competition. However, because the arena of sperm competition is within the female reproductive tract, thus the cryptic female choice determines the fate of sperm. How does the cryptic female choice dictate the rules of sperm competition? What are the underlying genes? In order to investigate the genetic basis of cryptic female choice and its role in conspecific sperm precedence, I perform quantitative trait loci analysis between Drosophila simulans and D. mauritiana. Two significant QTLs on X chromosome are detected that associated with BC1 (backcross generation one) female fecundity and may involve hybrid female sterility genes. One significant and one marginal significant QTLs on third chromosome are associated with cryptic female choice. Focusing on genes within these two intervals might be a good first step in approaching the basis of cryptic female choice. Potential candidate linked to the intervals may be olfactory receptors. Olfactory receptor may express on the sperm, ovary and/or female reproductive tract that could response to chemicals and/or hormones and allow female to bias the paternity. By comparing previous genetic studies, it also indicates the overlapping between QTLs of male traits and female mate choice. Linkage disequilibrium between male traits and female mate choice and female mate choice driven by postcopulatory process will be an interesting topic and could be further revealed in the two QTLs on the third chromosome.

Abstract    i
List of Figures    vii
Introduction    1
Sexual selection and sperm competition    1
Male strategies to conquer competitors in the sperm competition game    3
Male strategies to avoid direct sperm competition    5
Estimation of sperm competition    7
Cryptic female choice    8
Conspecific sperm precedence    9
Molecular mechanism of sperm competition and cryptic female choice    10
Conspecific sperm precedence between D. simulans and D. mauritiana    11
Materials and methods    13
Drosophila stocks    13
Sperm competition experiments    15
Paternity identification and estimation of Psim’    17
Genotyping    17
QTL analysis    19
Results    25
Overview of phenotypic variation among F1 females    25
Mating frequency in F1 and BC1 females    27
Overview of phenotypic variation in BC1, backcross generation one, females    27
Single marker analysis    29
QTL, Quantitative trait loci, analysis    31
Discussion    37
The sperm preference of F1 females is toward D. simulans males    37
Cryptic female choice in BC1 females    37
Composite interval mapping of fecundity    38
Comparison between genetic studies    40
Candidate genes of cryptic female choice    45
References    47
Appendix: raw data    53

List of Tables
Table 1. Single and double mating cross types    16
Table 2. Molecular and phenotypic markers    20
Table 3. Summary of single and double mating experiments    27
Table 4. Percentage of variance in Psim’, FBC1 and F2d among BC1females explained by the marker loci in single-marker analysis    30
Table 5. Predicted quantitative trait loci by composite interval mapping.    31
Table 6. Genetic basis of fecundity and female mate choice in Drosophila.    42
Table 7. Genetic basis of postcopulatory male traits and female preferences in Drosophila    43


List of Figures
Fig. 1. Generation of parental D. mauritiana (sn; j cn bw; irr) strain, F1 and BC1 females    14
Fig. 2. Time course of sperm competition experiments    16
Fig. 3. Genetic position of molecular markers    18
Fig. 4. The DNA electrophoresis photograph of molecular markers    24
Fig. 5. Numbers of progeny of F1 females after single or double matings    26
Fig. 6. The distribution of fecundity and Psim’ of FSM females    26
Fig. 7. Distribution of (A) F2d and (B) FBC1    28
Fig. 8. Distribution of sperm competitive success of D. simulans males (Psim’)    29
Fig. 9. Composite interval mapping for F2d    33
Fig. 10.  Composite interval mapping for FBC1    34
Fig. 11. Composite interval mapping for Psim’    35
Fig. 12. The overview of QTL analysis of F2d, FBC1 and Psim’    36
Fig. 13. Genetic position of markers in D. melanogaster against D. simulans genome.    39
Fig. 14. Scheme of genetic basis of pre- and postcopulatory male traits and female preferences    44

                                

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