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1、StagesofEmbryonicDevelopmentoftheZebrafish 刘永飞20121902004 斑马鱼 Daniorerio 属于辐鳍亚纲 Actinop terygii 鲤科 Cyprinidae 短担尼鱼属 Danio 的一种硬骨鱼 原产于印度东部 巴基斯坦 缅甸及孟加拉国的小溪 稻田及恒河中游地区 是一种常见的热带观赏鱼 因其体侧具有像斑马一样纵向的暗蓝与银色相间的条纹而得名 斑马鱼的发展史 一 20世纪70年代初 利用T4噬菌体研究遗传学的美国俄勒冈大学著名遗传学家GeorgeStreisinger注意到斑马鱼的优点 从宠物店购买了斑马鱼 开始研究其养殖方法 观察其
2、胚胎发育过程 发展一些相关的遗传学技术 二 近十年后 他的研究组于1981年在Nature上发表了第一篇具有深刻影响的论文 Productionofclonesofhomozygousdiploidzebrafish 在这篇论文中 他们报导了斑马鱼的体外受精技术 单倍体诱导技术 建立了纯合品系 并介绍了斑马鱼的第一个自然突变体 golden 三 在其后的几年中 Streisinger和他的同事先后报导了斑马鱼的卵裂特点 不同时期胚胎中细胞的发育命运等 并发现斑马鱼脑中的许多神经元的排列简单而有规律 这些研究成果的报导 证明了斑马鱼适合用做模式动物 引起了许多发育生物学家的关注 四 20世纪90
3、年代初 曾在果蝇上通过诱变研究阐明果蝇胚胎早期发育机理的德国发育生物学家ChristineN sslein Volhard 以及美国哈佛大学WolfgangDrie ver博士的研究组同时开始对斑马鱼进行大规模化学诱变研究 到1996年他们共鉴定出约4000种斑马鱼突变体 为研究脊椎动物发育的分子机理储备了丰富的遗传资源 Development专门于1996年用一期增刊发表了36篇相关论文 斑马鱼的特点 成年鱼个体小 易于饲养发育快速 性成熟期短 繁殖力强胚胎体外发育 易于观察和操作较完善的胚胎和遗传学操作技术基因组序列的注释已经完成品系资源丰富 StagesofEmbryonicDevelo
4、pmentoftheZebrafish 0 75 2 25h 0 0 75h 2 25 5 25h 5 25 10h 10 24h 24 28h 48 72h One ZygotePeriod 0 0 75h 1 Thenewlyfertilizedeggisinthezygotep erioduntilthefirstcleavageoccurs about40minutesafterfertilization 2 Thezygoteisabout0 7mmindiameteratthetimeoffertilization 3 Weincludeonlyasinglestagecalled
5、One cellstage howevermanychangesareoccur ring andonecouldeasilysubdividetheperi od StagesDuringtheZygotePeriod Thezygoteperiod A afewminutesafterfertilization B about10minafterfertil ization One cellstage Two CleavagePeriod 0 75 2 25h 1 Afterthefirstcleavagethecells divideatab out15 minuteintervals
6、2 Thecytoplasmicdivisionsaremeroblastic t heyonlyincompletelyundercuttheblastodisc andtheblastomeres oraspecificsubsetofthemaccordingtothestageremaininterconnectedbycytoplasmicbridges 3 Thesixcleavagesthatcomprisethisperiodf requentlyoccuratregularorientationssothato necanseehowmanyblastomeresarepre
7、senta rebytheirarrangement countingthemisunnec cessary StagesDuringtheCleavagePeriod Four cellstage 1h Eight cellstage 1 25h Sixteen cellstage 1 5h Thirtytwo cellstage 1 75h Sixtyfour cellstage 2h Two cellstage 0 75h Embryosduringthecleavageperiod A Two cellstage 0 75h 8 Four cellstage 1h C Eight ce
8、llstage 1 25h D Sixteen cellstage 1 5h E Thirty twocellstage 1 75h F Sixty fourcellstage 2h Three BlastulaPeriod 2 25 5 25h Weusethetermblastulatorefertot heperiodwhentheblastodiscbeginstolookball like atthe128 cellstage ore ighthzygoticcellcycle anduntiltheti meofonsetofgastrulation ca cycle14 Stag
9、esDuringtheBlastulaPeriod 128 cellstage 2 25h 256 cellstage 2 5h 512 cellstage 2 75h Highstage 3 25h lk cellstage 3h Oblongstage 3 66h Spherestage 4h Domestage 4 33h 30 epibolystage 4 66h Faceviewsofembryosduringtheblastulaperiod A 256 cellstage 2 5h B Highstage 3 3h C Transitionbetweenthehighandobl
10、ongstages 3 5h D T ransitionbetweentheoblongandspherestages 3 8h E Domestage 4 3h F 30 epibolystage 4 7h Four GastrulaPeriod 5 25 10h Themorphogeneticcellmove mentsofinvolution convergence andextensionoccur producingtheprimarygermlayersandtheembr yonicaxis StagesDuringtheGastrulaPeriod 50 epibolysta
11、ge 5 25h Germ ringstage 5 66h Shieldstage 6h 75 epibolystage 8h 90 epibolystage 9h Budstage 10h Developmentduringthegastrulaperiod A 50 epibolystage 5 25h B Germringsta ge 5 7h D Shieldstage 6h H 75 epibolystage 8h J 90 epibolystage 9h L Budstage 10h Five SegmentationPeriod 10 24h 1 Awonderfulvariet
12、yofmorphogeneticmo vementsnowoccur thesomitesdevelop therudimentsoftheprimaryorgansbecomevisib le thetailbudbecomesmoreprominentandtheembryoelongates Thefirstcellsdifferent iatemorphologically andthefirstbodymovem entsappear 2 Thetailbudappearsatthecaudalendofthelengtheningaxisduringthewholetime Sta
13、gesDuringtheSegmentationPeriod 2 somitestage 10 7h 5 somitestage 11 7h 14 somitestage 16h 20 somitestage 19h 25 somitestage 21 5h A 2 somitestage 10 7h F 5 somitestage 11 7h I 14 somitestage 16h M 20 somitestage 19h N 25 somitestage 21 5h Six PharyngulaPeriod 24 48h Theembryoismostevidentlynowabilat
14、erallyorganizedcreature enterin gthepharyngulaperiodwithawellde velopednotochord andanewlycompl etedsetofsomitesthatextendtotheendofalongpost analtail StagesDuringthePharyngulaPeriod Prim 5stage 24h Prim 12stage 30h Prim 25stage 36h High pecstage 42h A Prim 5stage 24h B Prim 12stage 30h F Prim 25sta
15、ge 36h H High pecstage 42h Seven HatchingPeriod 48 72h 1 Thetimeofhatchingisnotusefulasastagingindexf orthezebrafish incontrasttosomeothertypesofembry os becauseindividualswithinasingledevelopingclutchhatchsporadicallyduringthewhole3rddayofdevelopm ent atstandardtemperature andoccasionallylater 2 Wh
16、etherornotanembryohashatched itsdevelopme ntprogresses hourbyhour andgenerallyindividualsth athavespontaneouslyhatchedarenotmoredevelopment allyadvancedthanonesremainingintheirchorions 3 Wearbitrarilycallthecreatures embryos untiltheendofthe3rdday andafterward larvae whethertheyhavehatchedornot StagesDuringtheHatchingPeriod Protruding mouthstage 72h Pec finstage 60h Long pecstagea 48h A Long pecstagea 48h C Pec finstage 60h E Protruding mouthstage 72h Thanks
