摘要
1栉孔扇贝(Chlamys farreri)♀×海湾扇贝(Argopecten irradians)♂异源三倍体诱导及细胞遗传学研究
受精过程荧光显微观察表明,选择充分促熟的亲贝,20℃水温下,远缘杂交可以顺利进行。精子入卵后可以激活卵子完成减数分裂,进而精卵原核融合,卵裂,早期胚胎能够正常发育。观察初步证实异源精子参与了后代遗传物质的组成,具备异源多倍体诱导操作的可行性。
以50%的受精卵排出第1极体为诱导时机,以60 mg/L 6-DMAP为诱导剂量,处理15min,平均三倍体诱导率可达88.56%,孵化率可达53.52%。孵化后幼虫生长缓慢,死亡严重。受精后14天壳长仅为148μm左右,远低于自交对照组(173μm)。14天幼虫存活率仅0.00067%。
GISH分析表明,诱导获得的三倍体胚胎其染色体组由2套母本染色体组和1套父本染色体组构成,确为异源三倍体。约2%的分裂相出现母本偏向性染色体转变。类型包括染色体片段,及整条染色体的遗传转变。表明异源基因组并不稳定,存在复杂的核-核、核-质相互作用,可能与幼虫不能成活有关。2虾夷扇贝(Patinopecten yessoensis)三倍体诱导与培育
尝试低渗法诱导虾夷扇贝三倍体,获得最优诱导条件:12℃水温,以50%的受精卵排出第1极体为处理时机,20‰的盐度条件处理受精卵20min,D形幼虫平均三倍体得率可达91.24%,平均孵化率达52.94%。
倍性变化趋势:大规模诱导群体在D形幼虫至2龄成贝期间,以D形幼虫期三倍体率最高,达50.06%。随后,三倍体率急剧下降,到4月龄降至12.07%。从4月龄到8月龄三倍体率下降趋于缓慢,至8月龄降至7.27%。8月龄至24月龄成贝期间三倍体率基本稳定。
人工诱导三倍体在4月龄至24月龄期间均未表现出生长优势。24月龄成贝,三倍体平均壳高、壳长、壳宽、总重分别比二倍体对照低8.59%、10.33%、8.79%和24.78%。
3虾夷扇贝三倍体(P.yessoensis)育性与四倍体培育三倍体性成熟期比对照晚1月左右。共检测三倍体个体60只。性腺存在不同程度的发育,总体水平低下。雄性先于雌性发育。雄性性腺多为乳白色,个别微泛黄,雌性多为橘红色,个别颜色较淡。性比基本正常,1龄期雌雄比约1:1.18,2龄期雌雄比约1:1.50。未发现雌雄同体个体。
雄性三倍体共催产29只,无一排精。电镜检测3只雄性性腺,发现存在大量精子细胞。精核开始浓缩,呈现不规则多边形。部分精子细胞线粒体嵴稀少,甚至呈空泡状。内质网稀少,未发现顶体形成,也未发现成熟精子。
雌性三倍体共催产23只,4只排卵。卵子平均直径86.65±2.53μm,比二倍体大10.79%,体积大35.99%。相对育性(产卵量)1龄为3%,2龄为4%。荧光显微观察表明,三倍体的卵子具有受精能力,与二倍体的精子受精后大多数能够类似正常的分裂和发育。约30%的卵子减数分裂异常,染色体行为不同步。3n♀x2n♂,子代主要为非整倍体,约70%幼虫倍性介于2n和3n之间,30%介于3n和5n之间。D形幼虫孵化率仅0.23%,至7日龄未发现存活幼虫。
以20‰的盐度条件处理3n♀×2n♂受精卵20min抑制第1极体释放,胚胎最高四倍体率达75.12%。幼虫发育异常迟缓,约受精后65h到达担轮期,120h到达D形期,孵化率平均仅0.09%。7日龄存活率仅0.07%。
4三倍体虾夷扇贝(P.yessoensis)基因组甲基化水平的MSAP分析虾夷扇贝三倍体13只二倍体20只,来源于同一诱导群体。9对引物MSAP扩增共得到7206条条带,其中甲基化条带1988条(27.6%),包括全甲基化1095条(15.2%),半甲基化893条(12.4%)。
主要结论为:(1)虾夷扇贝基因组甲基化水平较高。(2)个体间基因组甲基化水平差异较大。(3)三倍化导致虾夷扇贝基因组甲基化降低。(4)甲基化水平与性别有关。
平均总甲基化率:雌性高于雄性。(5)平均总甲基化率:二倍体雌性>二倍体雄性>三倍体雌性>三倍体雄性。(6)三倍化不利于虾夷扇贝雌性性状表达和性腺发育。
1 Induction and cytogenetic study of allotriploid scallop in Chlamys farreri♀×Argopecten irradians♂cross
Prior to the allotriploid induction, the feasibility of fertilization between Zhikong scallop Chlamys farreri eggs and Bay scallop Argopecten irradians sperm was verified by epifluorescence microscopy. In 20℃seawater, crosses between C. farreri♀and A. irradians♂can be easily made with mature parents.
Triploidy and incubation rates of D-stage larvae with an average of 88.56% and 53.52% respectively, were obtained by Meiosis II inhibition, with 60 mg/L 6-deimethylaminopurine (6-DMAP) treatment for 15 min durations when 50% zygotes released the first polar body. The consequent larvae grew very slowly, and reached 148um in shell length, far behind of the control of C. farreri×C farreri cross (173μm) at 14 days post fertilization (DPF). The resultant larvae encountered high mortality when reached 6 DPF, and larvae survival dropped to 0.00067% at 14DPF. Using Genomic In situ Hybridization (GISH), the consequent larvae were validated to be real interspecies hybrids from C. farreri X A. irradians cross. Maternally biased chromosome conversion, including fragment conversion and whole chromosome conversion, were revealed in about 2% of all metaphases. These results might indicate the instability of the hybrid genome, with complicated interactions between nuclear-nuclear or nuclear-cytoplasm, which might give rise to the inviable larvae.
2 Induction and growth performance of triploid yesso scallop (Patinopecten yessoensis)
Hypotonicity shock, a new method to induce triploidy was tested. The highest triploidy reached 91.2.4% in average with a mean incubation rate 52.94%, with 20‰salinity treatment for 20 min durations, when 50% zygotes released the first polar body.
A large-scale population of triploidy induction group was made to test the ploidy alteration and growth performance from D-stage to 24-month age, by the method mentioned above. Triploidy was found to drop significantly from D-stage (50.06%) to 4-month age (12.07%), and then slightly to 8-month age (7.27%). From 8 months on, the ploidy of the induction group changed little.
Prior to growth comparisons, ploidy of each individual used was conformed by flow cytometry. No growth advantage of triploid population was found during the cultivation from 4-month to 24-month age. Mean height, length and width of shell, and body weight of triploids were 8.59%, 10.33%, 8.79% and 24.78% lesser than diploids respectively.
3 Relative fecundity and tetraploid induction of triploid yesso scallop (P. yessoensis)
Triploids reached sexual maturity with apparent gonad development about 1 month later than diploids. Among the 60 triploids sampled, gonad developed worse with various degree. Gonad of males developed prior to female, with cream-white appearance in general, while female gonad possessed orange-red looking. Sex ratio in triploids seemed to be normal, with female to male 1 : 1.18 at 1-year age, and 1:1.50 at 2-year age, with no hermaphroditic individual found.
None of the 29 male triploids spawned in spite of artificially thermal stimulation. Gonad from 3 males was investigated by electron microscopy. Many spermatids were found in male gonads, with condensed chromatin of various shape, and lacked endoplasmic reticulum. Mitochondria in some spermatids possessed lacked developed cristae, some of which even turned to empty vesicles. No acrosome was found, nor did sperm.
Spawning was successfully induced in 4 (17.39%) of the 23 triploid females, with a relative fecundity estimated to be 3% of normal diploids at 1 year age, and 4% at 2 year age. Mean diameter of these eggs was 86.65±2.53μm, 10.79% longer than that of diploids, and 35.99% bigger for volume than that of diploids. Eggs produced by female triploids were fertilized with sperm from one diploid male to investigate meiotic maturation in the eggs and the resultant larvae. Most of the eggs seemed to complete meioses normally. Aberrant meioses were observed in about 30% of the eggs, with unsynchronization and lagging of chromosomes. Flow cytometric analysis of the blastula stage larvae from the 3n×2n cross showed the same pattern, with one aneuploid peak lying between 2n and 3n possessing about 70% of the total area, and the other peak falling between 3n and 5n possessing about 30% of the total area. Mean survival of the fertilized eggs to D-stage for 3n×2n crosses (0.23%) was significantly lower than that for controls (75.67%).At day 7, no survival was observed.
The highest mean tetraploidy reached 75.12%, in 3n♀×2n♂cross by 20‰salinity treatment for 20 min durations, to inhibit the first polar body. The resultant larvae developed extraordinary slowly, and reached trochophore stage at about 65 h, and D-shaped stage 120 h. Incubation rate of D stage larvae was only 0.09%, and survival to 7-day age was only 0.07%.
4 MSAP analysis on triploid yesso scallop (P. yessoensis) 13 triploids and 20 diploids used in MSAP analysis were from the same triploid induction group. Using 9 pairs of selective primers, a total of 7206 amplified loci were identified, of which, 1988 loci (27.6%) were methylated including 1095 full-methylation loci (15.2%), and 893 hemi-methylation loci (12.4%).
Main conclusions: (1) methylation level was high in yesso scallop. (2) methylation level varied bigger among individuals. (3) triploidy resulted in lower methylation level in yesso scallop. (4) methylation level may be related to gender of yesso scallop. (5) for mean methylation level: diploid female >diploid male >triploid female >triploid male (6) triploidy hindered the development of female gonad in some extent.
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