摘要
海湾扇贝自 1982 年引入我国以来,已成为我国北方海区重要的养殖贝类。其作为雌
雄同体型生物,在自然情况下,既能自体受精,也能异体受精。一方面自交可增加纯合基
因型的出现频率,从而增加隐性有害基因表现的机会,导致近交衰退;另一方面自交也可
以作为品种纯化的有效手段,而在动植物育种中有着广泛的应用。
本研究主要从这两方面入手,通过比较不同有效繁殖群体数对子代的影响,研究了作
为雌雄同体型动物的海湾扇贝的近交生物学效应。同时,也利用其可自交的特点,选择优
良个体进行连续自交培育了 F1、F2、F3代自交系;比较了自交系间的生长、存活等数量性
状,并估计了配合力等参数。另外,通过不同遗传背景的养殖群体间的混交,建立了海湾
扇贝的育种基础群体,并对其进行选择实验,估计了现实遗传力。还进行了海湾扇贝加拿
大群体和墨西哥湾扇贝的杂交实验,分析了 F1的杂种优势及母体效应。主要结果如下:
1. 采用不同有效繁殖群体数的海湾扇贝进行繁殖,研究了近交对海湾扇贝 F1、F2 的生
长、存活等生物学特性的影响。实验共设置了 6 个梯度,分别采用 1、2、10、30、50 个
亲贝进行繁殖,并以常规生产为对照组。在子一代,实验各组第 1 天的 D 形幼虫壳长并没
有显著差别(P>0.05)。随着幼虫的不断生长,差别越来越明显,表现为自交组的生长和存
活均低于其他各组, 而 Ne=2、10、30、50 及对照组间的差异并不显著(P>0.05)。养成期
间,自交组个体壳长、壳高、壳宽及总重的生长,均明显低于其他各组。另外,越冬期间,
自交组的存活率,也明显低于其他各组。在子二代,近交衰退不仅存在于 Ne=1 组,而且
也存在于 Ne=2 组。这两组的生长和存活都明显低于其他各组(Ne=10, 30, 50, control),
而后者之间没有显著区别。同时,Ne=1 组的近交衰退明显高于 Ne=2 组。由此可见,低的
有效繁殖群体数(Ne),增加了近交的可能性,能够对子代产生一些不利的影响。因此,在
苗种生产中维持较高的 Ne 水平是非常有必要的。而且,由于海湾扇贝较高的产生配子的
能力(单个亲贝就能够产生大量的配子),也可能导致近交水平的增加,因此推荐从不同的
养成地点选择亲贝进行苗种培育。
i
张海滨 海湾扇贝近交生物学效应和遗传改良研究
博士学位论文
2. 采用两个海湾扇贝的群体进行混合交配,建立了基础群体。经过一年的养殖,以壳
长为指标对所建立的基础群体进行选择,从 1300 个体中选择壳长最大的 10%个体(选择
强度为 1.755)进行繁殖,并随机从基础群体中选择相同数量的个体作为对照。结果显示,
在幼虫阶段选择组的生长和存活明显高于对照组(P<0.01), 现实遗传力为 0.52±0.24。由于
在稚贝期间,对照组的养殖密度相对较低,在养成阶段开始时(第 100 天),对照组的平
均壳长大于选择组。随着密度调整一致后,在养成阶段(第 100 天至 160 天)的生长速度
显著大于对照组。同时,选择组在这一阶段的存活率也明显高于对照组。实验说明,利用
不同遗传背景的海湾扇贝,建立遗传多样性高的基础群体,并在此基础上进行选择育种,
是一种较有潜力的海湾扇贝种质改良的方法。
3. 进行了海湾扇贝加拿大群体和墨西哥湾群体的杂交实验,对杂种一代及对照群体的
生长和存活进行了比较。四个子代群体的第一天的 D 形幼虫的壳长没有显著区别(94.27μm,
93.78μm, 93.48μm, 93.58μm)。在第 10 天,两个杂交组(CM, MC)的壳长(155.35μm,
156.14μm)显著大于两个对照群体(141.97μm, 146.20μm),且 MC 的生长速度大于 CM。同
时,存活率的杂种优势明显大于生长的杂种优势。在幼虫阶段,卵源和交配方式对壳长和
存活都具有显著的影响。杂种优势不仅存在于幼虫阶段,而且在稚贝及成体阶段一直存在。
在第 170 天,CM 的个体明显大于其它三个组合,虽然 MC 的壳长和壳高也大于两个对照
群体(MM,CC),但差异不显著。
4.从海湾扇贝群体中选择优良个体,建立了 F1、F2、F3 自交系,并研究各家系在幼
虫及养成阶段的生物学特征。实验结果表明,在不同的家系间存在着明显的遗传变异,因
而进行家系的选择和培育有着广泛的应用前景。经过三代的自交选育,已经得到一些生长
速度快且存活率高的家系。
5. 本研究中利用起源于F2自交系的三个杂交家系为材料,进行了3×3双列杂交,分析
各杂交组合的杂种优势,同时计算了各家系间的一般配合力和特殊配合力,为海湾扇贝杂
种优势的利用提供一定的理论依据。结果表明,31×40杂种优势最为明显,而且随着幼虫
的生长,杂种优势值呈增加趋势。杂种优势,一般配合力和特殊配合力存在一定的相关性,
一般配合力高的组合,特殊配合力也高,杂种优势也明显。
The Bay scallop, Argopecten irradians (Lamarck), has served as a primary cultured species
in north China since it was introduced from the United States in 1982 (Zhang FS et al., 1986).
This species is a functional hermaphrodite. It can self-fertilize and cross-fertilize in nature.
Self-fertilization can result in inbreeding depression by increasing the frequency of recessive
harmful gene. At the same time, self-fertilization can be seen as a useful tool to produce pure
lines and broadly used in breeding program.
In this study, inbreeding effects was evaluated by comparing growth and survival of F1, F2
with different effective population size. F1, F2, F3 families were obtained by self-fertilizing. The
growth and survival of families was observed, and combining ability were estimated. A base
population was produced by cross of two cultured populations of bay scallop, and the realized
heritability was estimated. A complete diallel cross between two bay scallop population,
Argopecten irradians concentricus Say (M) and Argopecten irradians irradians Lamarck (C),
was carried out, and heterosis and maternal effect were discussion.
The main results of this study are listed as follows:
1. The experiment was designed to study the effect of effective population size (Ne) on the
growth and survival of F1 and F2 bay scallop Argopecten irradians. To determine the magnitude
of these effects, six experimental groups were produced: Ne=1, 2, 10, 30, 50, control. At the first
generation, the shell length of Larvae for all experimental groups was not different from each
other at of 1th day D-larva. With the growth of the larvae, the growth and survival of Ne =1 group
were gradually smaller than that of the others, while there was no significant difference among
Ne=2, 10, 30, 50 and control groups. During grow-out stage, the growth of shell length, shell
iii
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
height, shell width and wet weight for Ne =1 group was lower than the others too. In winter,
survival rate of Ne=1 is lower than that of others (P <0.05). At second generation, inbreeding
depression exists not only in Ne=1 group but also in Ne=2 group. The growth and survival rates
of the two groups are significantly lower than those of the other groups (Ne=10, 30, 50, control).
There is no significant difference between the later (P>0.05). At the same time, the amount of
depression in Ne=1 group is significantly higher than that of Ne=2 group (P<0.05). These results
indicate that the low effective population size (Ne), which increases the possibility of inbreeding,
can lead to some harmful effects to the offspring. So it is essential to maintain a high level of Ne
in commercial seed production. Furthermore, as the high fecundity of bay scallop might lead to
increased inbreeding, selecting broodstock from different growout sites is recommended.
2. A base population was produced by cross of two cultured populations of bay scallop
Argopecten irradians irradians Lamarck. After 1 year rearing, truncation selection of the top
10% (i=1.755) was carried out in the base population of about 1300 adults. A control parental
group with identical number to the select parental group was randomly selected from the entire
population before isolation of the select parental group. The result showed that, at larval stage,
the growth rate of larvae in the selected line was significantly higher than that of control
(P<0.05), and that the realized heritability was estimated to be 0.52±0.24. Due to the lower
density of control at spat stage, mean shell length of control line was larger than that of select
line at day 110. With the density was adjusted to the same between two lines in grow out stage
(from day 110-160), daily growth rate of selected line was significantly higher than that of
control line (P<0.05). Survival of se
引文
常亚青, 刘小林, 相建海, 等. 2002. 栉孔扇贝中国种群与日本种群杂交子一代的早期生长
发育. 水产学报, 26(5): 385-390.
董双林, 王芳, 王俊, 等. 1999. 海湾扇贝对海水池塘浮游生物和水质的影响[J]. 海洋学报,
21(6): 138-144
郭平仲. 1993. 群体遗传学导论. 农业出版社. 158-233
黄佩民. 2002. 中国近代农作物育种事业发展史略(三)—提高创新阶段(1979~2000).
山西农业科学. 30 (4): 3-13.
姜景民, 孙海菁, 刘昭息.1996 火炬松纸浆材优良家系多性状选择 林业科学研究. 9(5);
455-460.
李新海, 袁力行, 李晓辉, 张世煌, 李明顺, 李文华, 2003. 利用 SSR 标记划分 70 份我国玉
米自交系的杂种优势群.中国农业科学. 36(6): 622-627
刘保忠, 张福绥, 何义朝. 1998. 几种神经活性物质对海湾扇贝幼虫变态诱导作用的研究.
海洋学报, 20(5): 55-60
刘小林, 常亚青, 相建海, 等,2003. 栉孔扇贝中国种群与日本种群杂交一代的中期生长发
育. 水产学报. 27(3): 193-199
刘晓, 张国范, 赵洪恩. 2003. 皱纹盘鲍“中国红”品系的选育. 38(4): 27
楼允东. 2001. 鱼类育种学. 中国农业出版社.1-106.
吕豪, 魏若飞. 1994. 太平洋牡蛎与大连湾牡蛎杂交实验. 水产科学, 13(6): 8-11
盛志廉, 陈瑶生. 1999. 数量遗传学. 科学出版社. 42-108
童一中等. 1979. 作物遗传育种知识. 上海: 上海科学技术出版社.
汪德耀, 刘汉英. 1959. 牡颇人工杂交的初步研究.动物学报, 11(3): 283-295.
杨爱国,王清印,刘志鸿,等. 2003. 虾夷扇贝×栉孔扇贝人工受精过程的荧光显微观察. 海
洋水产研究, 23(3): 1-4.
杨纪珂. 1979. 数量遗传基础知识. 北京: 科学出版社.
袁隆平, 2001. 我在杂交水稻方面所作的工作. 中国科技奖励. 9 (1): 14-19
张爱民, 刘冬成, 聂秀玲, 等. 2002. 杂种小麦育种的战略. 中国农业科技导报. 4(5): 42-48.
张福绥, 何义朝, 亓铃欣, 等. 1997. 海湾扇贝引种复壮研究. 海洋与湖沼, 28(2): 146-152
95
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
张福绥、何义朝、刘祥生等,1986,海湾扇贝引种育苗及试养,海洋与湖沼,17(5):367
—374
张福绥. 1993. 海湾扇贝引进中国 10 周年. 齐鲁渔业. (5): 9-12.
张国范, 1992. 中国近海栉孔扇贝遗传结构及遗传变异与生长的关系. 青岛: 中国科学院海
洋所.
张国范, 刘述锡, 刘晓, 等. 2003. 海湾扇贝自交家系的建立和自交效应. 中国水产科学,
10(6): 441-445
张国范, 王继红, 赵洪恩, 阙华勇, 刘晓, 2002. 皱纹盘鲍中国群体和日本群体的自交与杂
交 F1的 RAPD 标记. 海洋与湖沼 33(5): 484-491
张国范, 张福绥. 1993. 贝类遗传多样性及其永续利用(I) . 海洋科学. (5): 17-21
张国范,刘晓,阙华勇,等. 2004. 贝类杂交及杂种优势理论和技术研究进展. 海洋科学.28(7):
54-60.
张国范,薛真福. 1999. 我国养殖贝类大规模死亡的原因分析及防治对策. 中国水产. (9):
34-39
张劳, 李玉奎. 群体遗传学概论. 中国农业出版社, 1999.
张喜昌, 梁玉波, 刘仁沿, 等. 2002. 海湾扇贝养殖群体遗传多样性的研究. 海洋学报, 24(2):
107-113
赵洪恩, 1999. 鲍的增养殖. 沈阳: 沈阳出版社,
赵素君, 欧江涛, 白文林, 2002. 杂种优势遗传基础的研究进展. 西南民族学院学报自然科
学版, 38: 327-330
赵自仙, 高样扩, 杨克昌, 孔建华, 2002. 玉米自交系选育的方法综述. 种子.(3): 42-44.
郑怀平, 刘晓, 张国范, 等. 2004. 海湾扇贝杂交家系与[交家系生长和存活的比较. 水产学
报, 28(3): 267-272.
周茂德, 高允田, 吴融. 1982. 太平洋牡蛎与近江牡蛎、褶牡蛎人工杂文的初步研究. 水产
学报, 6(3):235-241.
庄杰云, 樊叶杨, 吴建立, 等. 2001. 超显性效应对水稻杂种优势的重要作用. 中国科学, C
辑, 31(2): 106-113
Allendorf F W, Phelps S R. 1980. Loss of genetic variation in a hatchery stock of cutthroat trout.
Trans. Am. Fishi. Soc., 109: 537-543.
Allendorf F.W. and Ryman, N., 1987. Genetic management of hatchery stocks. In: N. Ryman and
96
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
F. Utter (Editors), Population Genetics and Fishery Management, Washington Sea Grant
Program. University of Washington Press, Seattle, WA, pp.141-159
Allendorf, F.W. and Utter, F.M., 1979. Population genetics. In: W.S. Hoar, D.J. Randall and J.R.
Brett (Editors), Fish Physiology. Vol. VIII. Academic Press, New York, NY, pp. 407-454.
Apostolov, A, Slanev, S. 2002. Heterosis effect in different schemes of crossing and
hybridization in pig breeding. Zhivotnov'Dni Nauki. 39: 75-78.
Aulstad, D., Kittelsen, A., 1971. Abnormal body curvature of rainbow trout Salmo gairdneri
inbred fry. J. Fish. Res. Board Can. 28: 1918-1920
Barber, B.J. and N.J. Blake, 1991. Reproductive physiology. In, Scallops: Biology, Ecology and
Aquaculture, edited by S.E. Shumway, Elsevier Science, Amsterdam, pp.377-428.
Barber, B.J., Carnegie, R.B., Davis, C.V. and Mook, W., 1996. Effect of timing of seed
deployment on growth and mortality of oysters, Crassostrea virginica, affected by juvenile
oyster disease (JOD). J. World Aquacult. Soc. 27: 443–448
Barber, B.J., Davis, C.V. and Crosby, M., 1998. Cultured oysters, Crassostrea virginica,
genetically selected for fast growth also exhibit increased tolerance of Juvenile Oyster
Disease (JOD). J. Shellfish Res. 17: 1171–1175
Barrett S C H, Kohn J R., 1991. Genetic and evolutionary consequences of small population size
in plants: implications for conservation. Genetics and Conservation of Rare Plants (eds D A
Falk, K E Holsinger). pp: 3-30. Oxford University Press USA. New York.
Baud, J.P., Gérard, A. and Naciri-Graven, Y., 1997. Comparative growth and mortality of
Bonamia ostreae-resistant and wild flat oysters, Ostrea edulis, in an intensive system: I.
First year of experiment. Mar. Biol. 130: 71–79
Bayne, B. L., D. Hedgecock, D. J. Mcgoldrick, and R. Rees, 1999. Physiological differences
between inbred and hybrid Pacific oysters (Crassostrea gigas) support the efficiency
hypothesis of heterosis. J. Exp. Mar. Biol. Ecol. 233:115-130.
Beattie J H, Perdue J, Hershberger W, Chew K. 1987. Effects of inbreeding on growth in the
Pacific oyster (Crassostrea gigas). J. Shellfish Res., 6:25-28.
Beattie, J.H., Chew, K.K. and Hershberger, W.K., 1980. Differential survival of selected strains
of Pacific oysters (Crassostrea gigas) during summertime mortality. Proc. Natl. Shellfish.
Assoc. 70: 184–189
97
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
Beattie, J.H., Hershberger, W.K., Chew, K.K., Mahnken, C., Prentice, E.F., Jones, C., 1978.
Breeding for resistance to summertime mortality in the Pacific oyster (Crassostrea gigas).
Pub. No. WSG 78-3. Washington Sea Grant, Seattle, Washington, USA. 13 pp.
Beaumont A.R, Turner G., Wood A.R and Skibinski D.O. F, 2004. Hybridisations between
Mytilus edulis and Mytilus galloprovincialis and performance of pure species and hybrid
veliger larvae at different temperatures. J. Exp. Mar. Biol. Ecol. 302: 177-188.
Beaumont AR, 1991. Genetic studies of laboratory reared Mytilus edulis Heterozygote
deficiencies, heterozygosity and growth. Biol J Linn Soc 44: 273-285
Beaumont, A.R., Abdul-Matin, A.K.M. and Seed, R., 1993. Early development, survival and
growth in pure and hybrid larvae of Mytilus edulis and M. galloprovincialis. J. Molluscan
Stud. 59: 120–123.
Becker, W.A., 1984. Manual of Quantitative Genetics, 5th ed. Academic Enterprises, Pullman,
WA, USA.190 pp.
Bierne, N., David, P., Boudry, P. and Bonhomme, F., 2002. Assortative fertilization and selection
at larval stage in the mussels Mytilus edulis and M. galloprovincialis. Evolution 56:
292–298.
Bierne, N., Launey, S., Naciri-Graven, Y., Bonhomme, F., 1998. Early effect of inbreeding as
revealed by microsatellite analyses on Ostrea edulis larvae. Genetics 148: 1893–1906.
Bondari K. and Dunham R.A. 1987, Effects of inbreeding on economic traits of channel catfish.
Theor. Appl. Genet. 74: 1–9.
Bosworth B.G., Wolters W.R. and Saxton A.M., 1994. Analysis of a diallel cross to estimate
effects of crossing on performance of red swamp crawfish, Procambarus clarkii,
Aquaculture, 121: 301-312.
Bricelj, V.M., Ford, S.E., Borrero, F.J., Perkins, F.O., Rivara, G., Hillman, R.E., Elston, R.A.
and Chang, J., 1992. Unexplained mortalities of hatchery-reared, juvenile oysters
Crassostrea virginica (Gmelin). J. Shellfish Res. 11: 331–347
Bruce, A. B., 1910. The Mendelian theory of heredity and the augmentation of vigor. Science 32:
627-628
Castagna M, Duggan W. 1971. Rearing the bay scallop, Argopecten circularis, Proc. Natl.
Shellfish. Assoc., 61: 80-85.
98
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
Chiyokubo, T., Shikano, T., Nakajima, M., Fujio, Y., 1998. Genetic features of salinity tolerance
in wild and domestic guppies Poecilia reticulata . Aquaculture 167, 339–348.
Collins, G.N. 1921. Dominance and vigor of first generation hybrid. Am. Nat. 55: 116-133
Cotman, D.W. et.al. 1998. Birth weight and neonatal survival of harbour seal pops are positively
correlated with genetic variation measured by microsatellites. Proc. R. soc. London B Biol.
Sci. 265: 803-809
Cragg, S.M. and Crisp, D.J., 1991. The biology of scallop larvae. In, Scallops: Biology, Ecology
and Aquaculture, edited by S.E. Shumway, Elsevier Science, Amsterdam, pp. 75-132.
Crenshaw, Jr., J.W., Heffernan, P.B., Walker, R.L., 1991. Heritability of growth rate in the
southern bay scallop, Argopecten irradians concentricus (Say, 1822). J. Shellfish Res. 10:
55–63.
Crow J F, Dominance and overdominance. In: J W Gowen, Editor, Heterosis, Iowa State College
Press, Ames (1952), pp. 282–297.
Crow J F. Breeding structure of population, 1954, pages 543-556 In Kempthorne O, Bancroft T
A, Gowen J W et al. editors. Statistics and Mathematics in biology. The Iowa state college,
Iowa
Crow, J. F. 1998. 90 Years Ago: The Beginning of Hybrid Maize. Genetics 148: 923–928
Crow, J.F. 1948. Alternative hypothesis of hybrid vigor. Genetics 33: 478-487
Cruz P., Ibarra A. M., 1997. Larval growth and survival of two catarina scallop (Argopecten
circularis, Sowerby, 1835) populations and their reciprocal crosses. J. Exp. Mar. Biol. Ecol.
95-110.
Cruz, P., Ramirez, J.L., Garcia, G.A. and Ibarra, A.M., 1998. Genetic differences between two
populations of catarina scallop (Argopecten ventricosus) for adaptations for growth and
survival in a stressful environment. Aquaculture 166: 321–335
Daniels, S.J. and Walters. J.R. 2000. Inbreeding depression and its effects on natal dispersal in
red-cockaded woodpecker. Condor 102: 482-491
David H. Reed, Julian J. O'Grady, Barry W. Brook, Jonathan D. Ballou and Richard Frankham
Estimates of minimum viable population sizes for vertebrates and factors influencing those
estimates, Biological Conservation, In Press, Corrected Proof, Available online 6 February
2003,
99
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
David, P., B. Delay, P. Berthou, and P. Jarne, 1995. Alternative models for allozyme-associated
heterosis in the marine bivalve Spisula ovalis.. Genetics 139:1719-1726
David, P., Jarne, P., 1997. Heterozygosity and growth in the marine bivalve Spisula ovalis:
testing alternative hypotheses. Genet. Res. 70: 215–223.
Davidson, E.H., 1986. Gene Activity in Early Development. Academic Press, Orlando, FL.
Davis C.V.and Barber B.J. 1999.Growth and survival of selected lines of eastern oysters,
Crassostrea virginica (Gmelin 1791) affected by juvenile oyster disease. Aquaculture, 178:
253-271
Davis, C.V. and Barber, B.J., 1994. Size-dependent mortality in hatchery-reared populations of
oysters, Crassostrea virginica, Gmelin 1791, affected by juvenile oyster disease. J. Shellfish
Res. 13: 137–142
Davis, C.V., 2000. Estimation of narrow-sense heritability for larval and juvenile growth traits in
selected and unselected sub-lines of eastern oysters, Crassostrea virginica. J. Shellfish Res.
19: 613.
Davis, C.V., Hawes, R.O. and Hidu, H., 1991. Selective breeding of eastern oysters (Crassostrea
virginica) in a Maine estuary. J. Shellfish Res. 10: 294
De Vicente M C.1993. QTL analysis of transgressive segregation in an interspecific tomota cross.
Genetic, 134: 585—596
Dillon Jr., R.T., 1992. Minimal hybridisation between populations of the hard clams Mercenaria
mercenaria and Mercenaria campechiensis co-occurring in South Carolina. Bull. Mar. Sci.
50:. 411–416
Dillon, R.T.J. and Manzi, J.J., 1987. Hard clam, Mercenaria mercenaria, broodstocks: genetic
drift and loss of rare alleles without reduction in heterozygosity. Aquaculture 60: 99–105
East, E. M., 1908. Inbreeding in corn. Rep. Conn. Agric. Exp. Stn. pp. 419–428.
East, E. M., 1936. Heterosis. Genetics 21: 375–397.
Eknath A E, Doyle R W. Effective population size and rate of inbreeding in aquaculture of indian
major carps. Aquaculture, 1990, 85:293-305
Falconer, D.S., Mackay, T.F.C., 1996. Introduction to Quantitative Genetics. Longman Group,
Essex, England.
100
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
Farley, C.A. and Lewis, E.J., 1994. Studies of Juvenile Oyster Disease, 1993. J. Shellfish Res. 13:
314
Farley, C.A. and Lewis, E.J., 1995. Juvenile Oyster Disease studies 1994: epizootiology,
geographic occurrence. J. Shellfish Res. 14: 241
Farley, C.A., Lewis, E.J., Relyea, D. and Zahtila, J., 1995. Studies of resistance in progeny of
broodstock selected from Juvenile Oyster Disease (JOD) survivors. J. Shellfish Res. 14, p.
242
Farley, C.A., Lewis, E.J., Relyea, D., Zahtila, J. and Rivara, G., 1996. Resistance studies for
juvenile oyster disease (JOD). J. Shellfish Res. 15: 515
Fischer M and Matthies D., 1998. Effects of population size on performance in the rare plant
Gentianella germanica. J. Ecol.. 86:195-204
Ford, S.E. and Haskin, H.H., 1987. Infection and mortality patterns in strains of oysters
Crassostrea virginica selected for resistance to the parasite Haplosporidium nelsoni (MSX).
J. Parasitol. 73: 368–376
Ford, S.E., 1988. Host–parasite interactions in eastern oysters selected for resistance to
Haplosporidium nelsoni (MSX) disease: Survival mechanisms against a natural pathogen.
Am. Fish. Soc. Spec. Publ. 18: 206–224
Franklin I R. Evolutionary changes in small populations. 1980, Pages 135–150 in M. Soule′ and
B. A. Wil-cox, editors. Conservation biology: an evolutionary-ecological perspective.
Sinauer Associates, Sunderland, Massachusetts
Fujio, Y., 1982. A correlation of heterozygosity with growth rate in the Pacific oyster,
Crassostrea gigas. Tohoku J. Agric. Res., 33: 66-75.
Gaffney, P.M., Davis, C.V. and Hawes, R.O., 1992. Assessment of drift and selection in
hatchery populations of oysters (Crassostrea virginica). Aquaculture 105: 1–20.
Gaffney, P.M., Scott, T.M., Koehn, R.K. and Diehl, W.J., 1990. Interrelationships of
heterozygosity, growth rate and heterozygote deficiencies in the coot clam, Mulinia
lateralis. Genetics 124: 687–699
Gardner, J.P.A., Skibinski, D.O.F. and Bajdik, C.D., 1993. Shell growth and viability differences
between the marine mussels Mytilus edulis (L.), Mytilus galloprovincialis (Lmk.) and their
hybrids from two sympatric populations in S.W. England. Biol. Bull. 185: 405–416.
101
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
Gjedrem T., 1997. Selective breeding to improve aquaculture production. World Aquacult. 28:
33–45.
Gjedrem, T., 1983. Genetic variation in quantitative traits and selective breeding in fish and
shellfish. Aquaculture 33: 51–72
Gjerde B. 1993. Breeding and selection. In: Heen K, Monahan R L, Utter F (Eds.), Salmon
Aquaculture. Fishing News Book, Cambridge, MA, USA, pp. 187-208.
Gjerde et al., 1983. Gjerde, K. Gunnes and T. Gjedrem , Effect of inbreeding on survival and
growth in rainbow trout. Aquaculture 34: 327–332.
Gjerde, B., 1986. Growth and reproduction in fish and shellfish. Aquaculture 57: 37–55
Gjerde, B., Gunnes, K., Gjedrem, T., 1983. Effects of inbreeding on survival and growth in
rainbow trout. Aquaculture 34: 327–332.
Gosling, E.M., 1982. Genetic variability in hatchery produced Pacific oysters Crassostrea gigas
Thunberg). Aquaculture, 26: 273-287.
Griffing, 1956. Concept of general and specific combining ability in relation to diallel crossing
systems. Aust. J. Biol. Sci. 9: 463-493
Griffing, B., 1990. Use of a controlled-nutrient experiment to test heterosis hypotheses. Genetics,
126: 753-767.
Hadley, N.H., Dillon Jr., R.T., Manzi, J.J., 1991. Realized heritability of growth rate in the hard
clam Mercenaria mercenaria. Aquaculture 93: 109–119.
Haley, L.E., Newkirk, G.F., 1982. The genetics of growth rate of Crassostrea virginica and
Ostrea edulis. Malacologia 22: 399-401
Haley, L.E., Newkirk, G.F., Waugh, D.W., Doyle, R.W., 1975. A report on the quantitative
genetics of the American oyster. Crassostrea virginica under laboratory conditions. 10th
European Symposium on Marine Biology. Ostend. Belgium, September 17-23, 1: 221-228
Hans Bernhard Bentsenand Ingrid Olesen., Designing aquaculture mass selection programs to
avoid high inbreeding rates, Aquaculture, 204: 349-359
Haskin, H.H. and Ford, S.E., 1978. Mortality patterns and disease resistance in Delaware Bay
oysters. Proc. Natl. Shellfish. Assoc. 68, p. 80
Haskins, H.H., Ford, S.E., 1988. Characteristics of inbred oyster strains selected for resistance to
Haplosporidium nelsoni (MSX). J. Shellfish Res. 7, 162.
102
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
Hawes, R.O., K. Scully and H. Hidu, 1990. Growth rate of two diverse populations of American
oysters, Crassostrea virginica, and their reciprocal crosses. Aquaculture, Special Issue
Genetics in Aquaculture Ⅲ, pp.327.
Hawkins, A.J.S., Bayne, B.L., Day, A.J., et al., 1989. Genotype-dependent interrelations between
energy metabolism, protein metabolism and fitness. In, Reproduction, Genetics and
Distributions of Marine Organisms, edited by J.S. Ryland and P.A. Tyler, Olsen and Olsen,
Fredensborg, Denmark, pp. 283-292.
Hedgecock D, Chow V, Waples R S. 1992. Effective population numbers of shellfish
broodstocks estimated from temporal variance in allelic frequencies. Aquaculture, 108:
215-232.
Hedgecock, D., Cooper, K. and Hershberger, W., 1991. Genetic and environmental components
of variance in harvest body size among pedigreed Pacific oysters Crassostrea gigas from
controlled crosses. J. Shellfish Res. 10, p. 516.
Hedgecock, D., McGoldrick, D.J., Bayne, B.L., 1995. Hybrid vigor in Pacific oysters: an
experimental approach using crosses among inbred lines. Aquaculture 137: 285– 298.
Hedgecock, D., McGoldrick, D.J., Manahan, D.T., Vavra, J., Appelmans, N. and Bayne, B.L.,
1996. Quantitative and molecular genetic analyses of heterosis in bivalve molluscs. J. Exp.
Mar. Biol. Ecol. 203: 49–59.
Hedgecock, D., Sly, F.L., 1990. Genetic drift and effective population sizes of hatchery
propagated stocks of the Pacific oyster Crassostrea gigas. Aquaculture 88: 21– 38.
Heffernan, P.B., Walker, R.L. and Crenshaw Jr., J.W., 1992. Embryonic and larval responses to
selection for increased rate of growth in adult bay scallops, Argopecten irradians
concentricus (Say, 1822). J. Shellfish Res. 11 : 21–25
Heffernan, P.B., Walker, R.L. and Ryan, M., 1993. Second heritability estimate of growth rate in
the southern bay scallop, Argopecten irradians concentricus (Say, 1822). J. Shellfish Res.
12 p.151
Hershberger, W.K., Myers, J.M., Iwamoto, R.N., Meauley, W.C. and Saxton, A.M., 1990.
Genetic changes in the growth of coho salmon (Oncorhynchus kisutch) in marine net-pens,
produced by ten years of selection. Aquaculture 85: 187–197
Hershberger, W.K., Perdue, J.A., Beattie, J.H., 1984. Genetic selection and systematic breeding
103
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
in Pacific oyster culture. Aquaculture 39: 237– 245.
Hervio, D., Bachere, E., Boulo, V., Cochennec, N., Vuillemin, V., Le Coguic, Y., Cailletaux, G.,
Mazurie, J. and Mialhe, E., 1995. Establishment of an experimental infection protocol for
the flat oyster, Ostrea edulis, with intrahaemocytic protozoan parasite, Bonamia ostreae:
application in the selection of parasite-resistant oysters. Aquaculture 132: 183–194
Hilbish T.J., Winn E.P., Rawson P.D., 1993. Genetic-variation and covariation during larval and
juvenile growth in Mercenaria mercenaria. . Mar. Biol. 115: 97–104.
Houle D, 1994. Adaptive distance and the genetic basis of heterosis. Evolution 48: 1410–1417
HU, Y.P., R. A. Lutz, and R. C Vrijenhoek, 1993. Overdominance in early life stages of an
American oyster strain. J. Hered. 84:254-258.
Hussain M.G., Islam M.S., Hossain M.A., et al., 2002. Stock improvement of silver barb
(Barbodes gonionotus Bleeker) through several generations of genetic selection.
Aquaculture 204: 469-480.
Ibarra A.M., Ramirez J.L., Ruiz C.A., Cruz P., Avila S., 1999. Realized heritabilities and genetic
correlation after dual selection for total weight and shell width in catarina scallop
(Argopecten ventricosus). Aquaculture 175: 227-241.
Ibarra, A.M., Cruz, P. and Romero, B.A., 1995. Effects of inbreeding on growth and survival of
self-fertilized catarina scallop larvae, Argopecten circularis. Aquaculture 134: 37–47
Innes, D.J. and Haley, L.E., 1977. Genetic aspects of larval growth under reduced salinity in
Mytilus edulis. Biol. Bull. Mar. Biol. Lab. 153: 312–321
Jarayabhand, J. and Thavornyutikarn, M., 1995. Realized heritability estimation on growth rate
of oyster Saccostrea cuucllata Born, 1778. Aquaculture 138: 111–118.
Jones, D. F., 1917 Dominance of linked factors as a means of accounting for heterosis. Proc.
Natl. Acad. Sci. USA 3:310-312
Jones, D. F., 1918 The effects of inbreeding and crossbreeding upon development. Conn. Agric.
Exp. Stn. Bull. 107. 100 pp.
Jones, D. F., 1922 The productiveness of single and double first generation corn hybrids. J. Am.
Soc. Agron. 14: 242–252.
Jones, R., Bates, J.A., Innes, D.J., Thompson, R.J., 1996. Quantitative genetic analysis of growth
in larval scallops (Placopecten magellanicus). Mar. Biol. 124: 671–677.
104
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
Kacser. H. and J.A. Burns, 1981. The molecular basis of dominance. Genetics, 97: 639-666.
Keeble, F. and C. Pellew, 1910. The mode of inheritance of stature and of time of flowering in
peas (Pisum sativum). J. Genet. 1: 47-56
Keller, F. K and Waller. D. M, Inbreeding effects in wild populations. Trends Ecol Evol. 2002,
17: 230-241.
Keller, L.F. et al., Selection against inbred song sparrows during a natural population bottleneck.
Nature, 1994, 372: 356-357
Kincaid, H.L., 1976a. Effects of inbreeding on rainbow trout populations. Trans. Am. Fish. Soc.
105, 273–280.
Kincaid, H.L., 1976b. Inbreeding in rainbow trout Salmo Gairdneri . J. Fish. Res. Board Can. 33:
2420–2426.
Kincaid, H.L., 1977. Rotational line crossing: an approach to the reduction of inbreeding
accumulation in trout broodstocks. Prog. Gish-Cult. 39: 179-181
Kincaid, H.L., 1983. Inbreeding in fish populations used for aquaculture. Aquaculture 33:
215–227.
Koehn, R.K., 1991. The genetics and taxonomy of species in the genus Mytilus. Aquaculture, 94:
125-145
Kraeuter, J.N., L. Adamkewicz, M. Castagna, R. Wall and R. Karney, 1984. Rib number and
shell color in hybridized subspecies of the Atlantic Bay Scallop, Argopecten circularis.
Nautilus, pp. 17-20.
Lamkey, K.R and J. W. Edwards, 1998. Heterosis: theory and estimation. P. 62-77. Proceedings
34th Illinois Corn breeders’ School, Urbana, IL, 2-3 Mar. 1998. University of Illinois,
Urbana.
Lamkey, K.R. and J. W. Edwards, 1999. The quantitative genetics of heterosis. Pp. 31-38. In: JG
Coors and S. Pandey (ed.) Proceedings of the International Symposium on the Genetics and
Exploitation of Heterosis in Crops, CIMMYT, Mexico City, Mexico, 17-22 Aug. 1997.
Lande, R. 1995. Risk of population extinction from fixation of new deleterious mutations.
Evolution, 48: 1460-1469
Langdon C, Evans F, Jacobson D and Blouin M. 2003. Yields of cultured Pacific oysters
Crassostrea gigas Thunberg improved after one generation of selection. Aquaculture, 220:
105
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
227-244
Lannan J E. 1980. Broodstock mangement of Crassostrea gigas. I. Genetic variation in survival
in the larval rearing system. Aquaculture, 21: 323-336
Lannan, J.E., 1972. Estimating heritability and predicting response to selection for the Pacific
oyster, Crassostrea gigas. Proc. Natl. Shellfish. Assoc. 62: 62–66
Lannan, J.E., 1980. Broodstock management of Crassostrea gigas. IV. Inbreeding and larval
survival. Aquaculture, 21: 353-356.
Lannan, J.E., 1980. Broodstock management of Crassostrea gigas: III. Selective breeding for
improved larval survival. Aquaculture 21: 347–351.
Launey, S., Hedgecock, D., 2001. High genetic load in the Pacific oyster Crassostrea gigas.
Genetics 159: 255–265.
Lee, M., Taylor, G.T., Bricelj, V.M. and Ford, S.E., 1995. Continuing studies of the role of
bacteria (Vibrio spp.) in unexplained juvenile oyster mortalities. J. Shellfish Res. 14, p.243
Lee, M., Taylor, G.T., Bricelj, V.M., Ford, S.E. and Zahn, S., 1996. Evaluation of Vibrio spp.
and microplankton blooms as causative agents of juvenile oyster disease in Crassostrea
virginica (Gmelin). J. Shellfish Res. 15: 319–330
Lewis, E.J., Farley, C.A., Small, E.B. and Baya, A.M., 1996. A synopsis of juvenile oyster
disease (JOD) experimental studies in Crassostrea virginica. Aquat. Living Resour. 9:
169–178
Lewontin, R.C., 1974. The Genetic Basis of Evolutionary Change. Columbia University Press,
New York, NY, 346pp
Li Z.K., L. J. Luo, H. W. Mei, D. L. Wang, Q. Y. Shu, R. Tabien, D. B. Zhong, C. S. Ying, J. W.
Stansel, G. S. Khush, and A. H. Paterson, 2001. Overdominant Epistatic Loci Are the
Primary Genetic Basis of Inbreeding Depression and Heterosis in Rice. I. Biomass and
Grain Yield. Genetics, 158: 1737-1753
Losee, E., 1978. Influence of heredity on larval and spat growth in Crassostrea virginica. IN:
J.W. Avault (Editors), Proceedings of The Ninth Annual Meeting, World Mariculture
Society, pp.101-107.
Ludwig, A.N., 1989. Heritability, genetic correlation, and genotype-environment interaction of
larval and juvenile growth rate in the coot clam, Mulinia lateralis. J. Shellfish Res. 8 p 486.
106
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
Lukas F. Keller and Donald M. Waller., 2002. Inbreeding effects in wild populations, Trends in
Ecology & Evolution, 17: 230-241
Lutz C.G. and Wolters W.R. , 1989. Estimation of heritabilities for growth, body size, and
processing traits in red swamp crawfish, Procambarus clarkii (Girard). Aquaculture 78:
21–33.
Lymbery A.J., R.G. Doupe, G. Jenkins and T. Thorne, 2000. Genetics in the aquaculture industry.
Aquacult. Res. 31: 1–2.
Lynch, M., Conery, J. and Burger, R. 1995. Mutation accumulation and the extinction of small
populations. Am Nat, 146: 489-518
Madalena, F. E., 1993. A simple scheme to utilize heterosis in tropical dairy cattle. World Animal
Review, 74: 17-25.
Mallet, A.L. and Haley, L.E., 1983a. Effects of inbreeding on larval and spat performance in the
American oyster. Aquaculture, 33: 229-235.
Mallet, A.L., and L.E. Haley, 1983b. Growth rate and survival in pure population matings and
crosses of the oyster Crassostrea virginica. Can. J. Fish. Aquat.Sci., 57: 948-954.
Mallet, A.L., and L.E. Haley, 1984. General and specific combining abilities of larval and
juvenile growth and viability estimated from natural oyster populations. Mar. Biol., 81:
53-59
Manzi J.J., Hadley N.H., Dillion R.T., 1991. Hard clam, Mercenaria mercenaria, Broodstocks:
growth of selected hatchery stocks and their reciprocal crosses. Aquaculture, 94: 17-26
Mather, K and Jinks, J.L., 1971. Biometrical Genetics. Cornell University Press, Ithaca, NY,
USA, pp382.
Mcgoldrick, D. J., 1997 An experimental investigation of the genetic basis of heterosis in the
Pacific oyster Crassostrea gigas (Thunberg). Ph.D. Thesis, University of California, Davis,
CA.
McGoldrick, D.J. and Hedgecock, D., 1997. Fixation, segregation and linkage of allozyme loci
in inbred families of the Pacific oyster Crassostrea gigas (Thunberg): implications for the
causes of inbreeding depression. Genetics 146: 321–324.
Mitton JB and Grant MC, 1984. Associations among protein heterozygosity, growth rate and
developmental homeostasis. Annu Rev Ecol Syst 15: 479-499
107
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
Moav R. and Wohlfarth G., 1976. Two-way selection for growth rate in the common carp
(Cyprinus carpio L.). Genetics 82: 83–101.
Mrakovcic M, Haley L E. (1979) Inbreeding depression in the zebra fish Brachydanio rerio
(Hamilton Buchana). J. Fish Biol., 15: 323–327.
Nagler J.J., Parsons J.E and Cloud J.G., 2000. Single pair mating indicates maternal effects on
embryo survival in rainbow trout, Oncorhynchus mykiss. Aquaculture, 184: 177-183.
Nei M. Molecular population genetics and evolution[M]. North-Holland, Amsterdam and New
York, 1975, p88-89
Nei, M. and Roychoudhury, A.K. 1974. Sampling variances of heterozygosity and genetic
distance. Genetics, 76: 379-390.
Nei, M., 1983. Genetic polymorphism and the role of mutation in evolution. In: Nei, M., and
Koehn R.(Editors), Evolution of Genes and Proteins. Sinauer Associates, Sunderland, MA,
pp.165-190.
Nei, M., and Graur, D., 1984. Extent of protein polymorphism and the neutral mutation theory.
Evol. Biol., 17: 73-118
Nell, J.A., Sheridan, A.K., Smith, I.R., 1996. Progress in a Sydney rock oyster, Saccostrea
commercialis (Iredale and Roughley), breeding program. Aquaculture 144: 295– 302.
Nell, J.A., Smith, I.R., Sheridan, A.K., 1999. Third generation evaluation of Sydney rock oyster
Saccostrea commercialis (Iredale and Roughley) breeding lines. Aquaculture 170: 195–
203.
Neter, J., W. Wasserman and M. Kutner, 1985. Applied linear statistical models. Richard D. Irwin,
2nd. Edition, pp 1127
Newkirk G.F., 1980. Review of the genetics and the potential for selective breeding of
commercially important bivalves. Aquaculture, 19: 209-228
Newkirk G.F., 1986. Controlled mating of the European oyster, Ostrea edulis. Aquaculture,
57:111-116
Newkirk G.F., D.L. Waugh and L.E. Haley, 1977. Genetics of larval tolerance to reduced
salinities in two populations of oysters, Crassostrea virginica. J. Fish. Res. Board Can., 34:
383-387
Newkirk G.F., Haley L.E., Waugh D.L. and Doyle, R.W., 1977. Genetics of larvae and spat
108
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
growth rate in the oyster, Crassostrea virginica. Mar. Biol., 41: 49-52.
Newkirk, G.F. and Haley, L.E., 1982. Progress in selection for growth rate in the European
oyster Ostrea edulis. Mar. Ecol. Prog. Ser. 10: 77–79
Newkirk, G.F. and Haley, L.E., 1983. Selection for growth rate in the European oyster Ostrea
edulis: response of second generation groups. Aquaculture 33: 149–155.
Nicholas F.W.,1980. Size of population required for artificial selection. Genet. Res. 35 : 85–105.
Ouborg, J., van Treuren, R. and van Damme, J.M.M. 1991. Morphological variation and fitness
components in populations of varying size of Salvia pratensis L. & Scabiosa columbaria L.
Oecologia, 86: 359-367.
Pante M J R, Gjerde B, McMillan I. Inbreeding levels in selected populations of rainbow trout,
Oncorhynchus mykiss. Aquaculture, 2001, 192: 213-224.
Paynter, K.T., Dimichele, L., 1990. Growth of tray-cultured oysters (Crassostrea virginica
Gmelin) in Chesa-peake Bay. Aquaculture 87, 289– 297.
Pogson, G.H. and E. Zouros, 1994. Allozyme and RFLP heterozygosities as correlates of growth
rate in the scallop Placopecten magellanicus: A test of the associative overdominance
hypothesis, Genetics, 137: 221-231.
Pogson, G.H., 1991. Expression of overdominance for specific activity at the
phosphoglucomutase-2 locus in the Pacific oyster, Crassostrea gigas. Genetics, 128:
133-141.
Rawson, P.D. and Hilbish, T.J., 1990. Heritability of juvenile growth for the hard clam
Mercenaria mercenaria. Mar. Biol. 105: 426–429.
Refstie, T., 1990. Application of breeding schemes. Aquaculture 85: 163–169
Relyea, D., 1994. Modifying oyster hatchery management to avoid losses due to unexplained
mortality of juvenile oysters. J. Shellfish Res. 13, p 319
Robertson, F.W. and E.C.R. Reeve, 1952. Heterozygosity, environmental variation and heterosis.
Nature, 170: 296-298.
Rohlf, F.J. and R.R. Sokal, 1981. Statistical tables. W.H. Freeman and Company, New York, pp
219
Rye and I.L. Mao , 1998. Nonadditive genetic effects and inbreeding depression for body weight
in Atlantic salmon (Salmo salar L.). Livest. Prod. Sci. 57: 15–22.
109
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
Ryman, N., Stahl, G., 1980. Genetic changes in hatchery stocks of brown trout Salmo trutta . Can.
J. Aquat. Sci. 37: 82–87.
Saccheri I, Kuussaari M, Kankare M, Vikman P, Fortelius W, Hanski L.1998. Inbreeding and
extinction in a butterfly population. Nature, 392: 491–494
Sarver, S.K., M. Katoh and D.W. Foltz, 1992. Apparent overdominance of enzyme activity in
two marine bivalves. Genetica, 85: 23 l-239.
Sastry A N. 1963. Reproduction of the bay scallop, Argopecten irradians Lamarck. Influence of
temperature on maturation and spawning. Biol. Bull. 125:146-153
Sbordont S, Matthaeis E D, Sbordont M C, et al. 1986. Bottleneck effects and the dipression of
genetic variability in hatchery stocks of Penaeus japonicus (Crustacea, Decapoda).
Aquacultre, 57:239-251
Sheridan A.K., 1997. Genetic improvement of oyster production-a critique. Aquaculture. 153:
165-179
Sheridan, A.K., Smith, I.R. and Nell, J.A., 1996. Reducing the impact of environmental variation
in a growth rate improvement program for the Sydney rock oyster Saccostrea commercialis.
Aquaculture 143: 145–154
Shikano T, Chiyokubo T, Taniguchi N. 2001. Effect of inbreeding on salinity tolerance in the
guppy (Poecilia reticulata). Aquaculture, 202: 45-55.
Shikano T. and Taniguchi, N. 2002. Using microsatellite and RAPD markers to estimate the
amount of heterosis in various strain combinations in the guppy (Poecilia reticulata) as a
fish model, Aquaculture, 204: 271-281
Shikano, T., Nakadate, M., Nakajima, M., Fujio, Y., 1997. Heterosis and maternal effects in
salinity tolerance of the guppy Poecilia reticulata. Fish. Sci. 63: 893–896.
Shull, G. H., 1908 The composition of a field of maize. Am. Breeders Assoc. Rep. 4: 296–301.
Shull, G. H., 1909. A pure line method of corn breeding. Am. Breeders Assoc. Rep. 5: 51–59.
Shull, G. H., 1914. Duplicated genes for capsule form in Bursa bursapastoris. Z. Indukt.
Abstammungs u. Vererbungsl. 12: 97–149.
Shull, G. H., 1948 What is “heterosis”?. Genetics 33: 439–446.
Singh, S.M. and Zouros, E., 1981. Genetics of growth rate in oysters and its implication for
aquaculture. Can. J. Genet. Cytol. 23: 119–130
110
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
Small, E.B., 1995. Ciliated protists associated with Juvenile Oyster Disease. J. Shellfish Res. 14:
247–248
Smith, I.R., Sheridan, A.K. and Nell, J.A., 1995. Evaluation of growing methods for use in a
Sydney rock oyster Saccostrea commercialis (Iredale and Roughley) selective breeding
program. Aquaculture 131: 189–195
Soule′ M E. Thresholds for survival: maintaining fitness and evolutionary potential. 1980, Pages
151–169 in M. E. Soule′ and B. A. Wilcox, editors. Conservation biology: an
evolutionary-ecological perspective. Sinauer, Sunderland, Massachusetts
Sprague, G.F. and Tatum, L.A. 1942. General vs. specific combining ability in single crosses of
corn. J. Amer, Soc. Agron., 34: 923-932
Stroemgren, T., Nielsen, M.V., 1989. Heritability of growth in larvae and juveniles of Mytilus
edulis. Aquaculture 80: 1–6.
Stuber, C. W., 1994. Heterosis in plant breeding. Plant Breed. Rev. 12: 227-251
Stuber, C.W., S.E. Lincoln, D.W. Wolff, T. Helentjaris and E.S. Lander, 1992. Identification of
genetic factors contributing to heterosis in a hybrid from two elite maize inbred lines using
molecular markers. Genetics, 132: 823-839.
Stuber. C.W. 1992. Biochemical and molecular marders in plant breeding. Plant Breeding Rev.,
9: 37-61
Su G S, Liljedahl L E, Gall G A E. 1996. Effects of inbreeding on growth and reproductive traits
in rainbow trout (Oncorhynchus mykiss). Aquaculture, 142:139-148
Takahito Shikano, Takashi Chiyokubo and Nobuhiko Taniguchi., Effect of inbreeding on salinity
tolerance in the guppy (Poecilia reticulata), Aquaculture, 202: 45-55
Taniguchi N, Sumantadinata K, Iyama S. (1983) Genetic change in the first and second
generations of hatchery stock of black sea bream. Aquaculture, 35: 309-320.
Tave, D., 1993. Genetics for Fish Hatchery Managers, 2nd edn. Van Rostrand-Reinhold, New
York, pp 415.
Toro, J.E. and Newkirk, G.F., 1991. Response to artificial selection and realised heritability
estimate for shell height in the Chilean oyster Ostrea chilensis. Aquat. Living Resour. 4:
101–108
Toro, J.E., Aguila, P. and Vergara, A.M., 1996. Spatial variation in response to selection for live
111
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
weight and shell length from data on individually tagged Chilean native oysters (Ostrea
chilensis Philippi, 1845). Aquaculture 146: 27–36.
Toro, J.E., Aguila, P., Vergara, A.M. and Newkirk, G.F., 1994. Realised heritability estimates
for growth from data on tagged Chilean native oyster (Ostrea chilensis). World Aquac. 25:
29–30.
Toro, J.E., Newkirk, G.F., 1990. Divergent selection for growth rate in the European oyster
Ostrea edulis: response to selection and estimate of genetic parameters. Mar. Ecol. Prog.
Ser. 62: 219–227.
Toro, J.E., Paredes, L.I., 1996. Heritability estimates of larval shell length in the Chilean blue
mussel Mytilus chilensis, under different food densities. Aquat. Living Resourc. 9: 347–350.
Toro, J.E., Sanhueza, M.A., Winter, J.E., Aguila, P., Vergara, M., 1995. Selection response and
heritability estimates for growth in the Chilean oyster Ostrea chilensis. J. Shellfish Res. 14:
87– 92.
Van Treuren, R., Bijlsma, R., van Delden, W. and Ouborg, N.J. 1991. The significance of
genetic erosion in the process of extinction. 1. Genetic differentiation in Salvia pratensis
and Scabiosa columbaria in relation to population size. Heredity, 66: 181-189
Van Vleck, L.D., E.J. Pollak and E.A.B. Oltenacu, 1987. Genetics for the animal sciences. W.H.
Freeman and Company, New York, pp 391.
Vrijenhoak, R.C., Ford, S.E. and Haskin, H.H., 1990. Maintenance of heterozygosity during
selective breeding of oysters for resistance to MSX disease. J. Hered. 81: 418–423
Wada, K., 1986. Genetic selection for shell traits in the Japanese pearl oyster, Pinctada fucata
martensii. Aquaculture 57: 171–176
Watterson, G.A., 1984. Allelic frequencies after a bottleneck. Theor. Popul. Biol., 26: 387-407.
Willhelm, R. and Hilbish, T.J., 1998. Assessment of natural selection in a hybrid population of
mussels: evaluation of exogenous vs. endogenous selection models. Mar. Biol. 131:
505–514.
Wright S. 1931. Evolution in mendelan populations. Genetics, 16: 97-159
Xiao J, Li J, Yuan L and Tanksley S D, 1995. Dominance is the major genetic basis of heterosis
in rice as revealed by QTL analysis using molecular markers. Genetics 135:876-879
Zhang Guofan, Liu Shuxi, Liu Xiao, Guo Ximing and Zhang Fusui. 2003. Self-fertilization
112
张海滨 海湾扇贝近交生物学效应和遗传改良研究 博士学位论文
family establishment in bay scallop Argopecten irradians. Journal of Fisheries Sciences of
China, 10: 441-445.
Zhang, F. S., Y. C. He, L. X. Qi, L. N. Sun and B.Z. Liu, 1994. Introduction and spat rearing of
the F1 generation of Argopecten irradians concentricus Say. Oceanologia et Limnologia
Sinica 25: 372 – 377 (in Chinese with English abstract).
Zheng H.P., Zhang G.F., Liu X., Zhang F., Guo X., 2004. Different responses to selection in two
stocks of bay scallop, Argopecten irradians irradians Lamarck (1819). J. Exp. Mar. Biol.
Ecol. 313: 213-223.
Zouros, E. and D.W. Foltz, 1984. Possible explanations of heterozygote deficiency in bivalve
molluscs. Malacologia, 25: 583-591.
Zouros, E. and D.W. Foltz, 1987. The use of allelic isozyme variation for the study of heterosis.
Isozymes: Curr. Top. Biol. Med. Res., 13: l-59.
Zouros, E. and Pogson, GH. 1994. Heterozygosity, heterosis and adaptation. In: Genetics and
Evolution of Aquatic Organisms, edited by A.R. Beaumont, Chapman and Hall, London,
pp.135-146
Zouros, E., 1987. On the relation between heterozygosity and heterosis: an evaluation of the
evidence from marine mollusks, Isozymes: Curr. Top. Biol. Med. Res., 15: 255-279.
113