INTESTINE OF ZEBRAFISH: REGIONALIZATION, CHARACTERIZATION AND STEM CELLS WANG ZHENGYUAN (M.Sci., NUS) (B.Eng., NPU) A THESIS SUBMITTED FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN COMPUTATION AND SYSTEMS BIOLOGY (CSB) SINGAPORE-MIT ALLIANCE NATIONAL UNIVERSITY OF SINGAPORE 2010 Acknowledgements I want to thank my supervisors, professor Matsudaira Paul and professor Gong Zhiyuan, whose time, knowledge and wise guidance has constituted a key component ensuring the continual progression of my research work. I want to thank my thesis committee members, professor Lodish Harvey and assistant professor Bhowmic Sourav, for following my progress, evaluating and steering my work. Thanks also go to professor Rajagopal Gunaretnam, who initiated the project and helped much to get the project started several years ago. This study was supported by funding and a graduate fellowship from the Singapore-MIT Alliance. Special thanks to my labmates, Zhan Huiqing, Wu Yilian, Du Jianguo, Tavakoli Sahar, Li Zhen, Zheng Weiling, Tina Sim Huey Fen, Liang Bing, Ung Choon Yong, Lam Siew Hong, Yin Ao, Mintzu, Li Caixia, Grace Ng, Sun Lili, Cecilia, Lana and others. Four years of doctoral study in the laboratory would by no means be joyful and fun-filled without the company of them. Last but not least, thanks go to my family members, including my parents, sisters, wife and son for their full support during my doctoral study. Time spent in the laboratory could not be spent with my family. Thank them for i bearing with me during the years of scientific training. Thank my son, the little lovely creature, for bringing oceans of joy to me. ii Contents Introduction 1.1 Introduction to the digestive system . . . . . . . . . . . . . . . . 1.2 Tissue architecture and cell types of the intestinal epithelium . . 1.3 Turnover of the intestinal epithelium . . . . . . . . . . . . . . . 1.4 Significance of the study of the digestive system . . . . . . . . . 1.5 Intestinal stem cells . . . . . . . . . . . . . . . . . . . . . . . . . 1.5.1 Location of intestinal stem cells . . . . . . . . . . . . . . 1.5.2 Intestinal stem cell number . . . . . . . . . . . . . . . . 1.5.3 Intestinal stem cell marker . . . . . . . . . . . . . . . . . 1.6 Intestines of different vertebrate models . . . . . . . . . . . . . . 1.6.1 Mouse intestine . . . . . . . . . . . . . . . . . . . . . . . 1.6.2 Chicken intestine . . . . . . . . . . . . . . . . . . . . . . 1.6.3 Frog intestine . . . . . . . . . . . . . . . . . . . . . . . . 1.6.4 Zebrafish intestine . . . . . . . . . . . . . . . . . . . . . 1.7 Establishing zebrafish as a vertebrate model for study on intestine 1.8 Research goals of the current work . . . . . . . . . . . . . . . . 1.8.1 Morphological and histological features of zebrafish intestine . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.8.2 Characterization of regionalization of zebrafish intestine through genome-wide gene expression analysis . . . . . . 1.8.3 Study of the cell fate decision in zebrafish intestine . . . 1.8.4 Responsive nature of intestine during regeneration . . . . 1.8.5 Computational analysis of intestinal stem cells and their adaptive changes . . . . . . . . . . . . . . . . . . . . . . 11 12 13 13 14 16 17 19 20 21 21 22 22 23 Functional organization along the rostrocaudal axis of the intestine 24 2.1 Background . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 i 2.2 2.3 Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . 2.2.1 Maintenance of zebrafish and dissection of zebrafish intestine . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.2.2 Paraffin sectioning of zebrafish intestine . . . . . . . . . 2.2.3 Hematoxylin and eosin and alcian blue staining . . . . . 2.2.4 Quantitative real-time PCR (qRT-PCR) . . . . . . . . . 2.2.5 Microarray experiments . . . . . . . . . . . . . . . . . . 2.2.6 Identification of differentially expressed genes from the microarray data . . . . . . . . . . . . . . . . . . . . . . . 2.2.7 Gene ontology (GO) analysis by GO Tree Machine . . . 2.2.8 Pathway analysis using WebGestalt . . . . . . . . . . . . 2.2.9 Gene Set Enrichment Analysis . . . . . . . . . . . . . . . Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.3.1 Architectural differences along the zebrafish intestinal tract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.3.2 Distinct molecular signatures along the zebrafish intestinal tract . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.3.3 Molecular features of the small and large intestine-like functions . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.3.4 Analysis of gene ontology along the anterior-posterior axis 2.3.5 Cross-species Gene Set Enrichment Analysis (GSEA) indicates the segments S1-S5 to be multi-functional . . . . 2.3.6 Stomach-like functions of the intestine . . . . . . . . . . 2.3.7 Pathway analysis for zebrafish intestine . . . . . . . . . . 2.3.8 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . 2.3.9 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . Regulation of cell fate and composition of the intestinal epithelium 3.1 Background . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2 Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . 3.2.1 DAPT treatment of zebrafish . . . . . . . . . . . . . . . 3.2.2 Alcian blue and Periodic Acid in Schiff’s reagent staining 3.2.3 Whole mount in situ hybridization . . . . . . . . . . . . 3.2.4 Cryosection of zebrafish intestine . . . . . . . . . . . . . 3.2.5 Immunohistochemistry . . . . . . . . . . . . . . . . . . . 3.3 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.3.1 Inhibition of Notch signaling in larval zebrafish intestine 27 27 27 28 28 29 30 30 31 32 32 32 37 43 48 53 56 59 66 69 70 71 72 72 73 74 74 75 76 76 ii 3.3.2 3.4 3.5 Verification of inhibition of Notch signaling in adult zebrafish intestine . . . . . . . . . . . . . . . . . . . . . . 3.3.3 Reduction in the pool of intestinal progenitor cells upon inhibition of Notch . . . . . . . . . . . . . . . . . . . . 3.3.4 Increase of secretory lineages after inhibition of Notch signaling . . . . . . . . . . . . . . . . . . . . . . . . . . 3.3.5 Enhanced expression of gata6 upon inhibition of Notch in the intestine . . . . . . . . . . . . . . . . . . . . . . 3.3.6 Enhanced activity of BMP signaling due to inhibition of Notch signaling . . . . . . . . . . . . . . . . . . . . . . 3.3.7 Suppression of glycogen-rich intestinal subepithelial myofibroblasts (ISEMFs) along the villus axis due to inhibition of Notch signaling . . . . . . . . . . . . . . . . . Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.4.1 Notch signaling and binary lineage allocation . . . . . . 3.4.2 Involvement of a distinct cohort of glycogen-rich ISEMFs in cell lineage allocation . . . . . . . . . . . . . . . . . 3.4.3 Preferable targeting of secretory cells in cancer . . . . . 3.4.4 Cooperative BMP and gata6 activities in epithelial differentiation . . . . . . . . . . . . . . . . . . . . . . . . Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77 . 78 . 81 . 84 . 85 . 89 . 93 . 93 . 95 . 97 . 98 . 100 Regeneration of zebrafish intestine following whole body gammaradiation 101 4.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102 4.2 Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105 4.2.1 Experiment setup for radiation . . . . . . . . . . . . . . 105 4.2.2 Sampling schedule . . . . . . . . . . . . . . . . . . . . . 105 4.2.3 RNA extraction and real-time PCR . . . . . . . . . . . . 106 4.2.4 Paraffin embedding and AB-PAS staining . . . . . . . . 106 4.2.5 Alkaline phosphatase staining . . . . . . . . . . . . . . . 106 4.3 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110 4.3.1 Survival of zebrafish after whole body gamma radiation . 110 4.3.2 Two rounds of elimination of intestinal villi . . . . . . . 110 4.3.3 Two waves of Wnt/beta-catenin signaling: a driver of proliferation . . . . . . . . . . . . . . . . . . . . . . . . . 113 4.3.4 Cell proliferation as measured by pcna staining . . . . . 116 4.3.5 Radiation induced cell apoptosis . . . . . . . . . . . . . . 118 4.3.6 Changes in the intestinal epithelium renewal . . . . . . . 122 iii 4.3.7 4.3.8 4.4 Regeneration of the secretory epithelial cells . . . . . . Maintenance of basic intestinal functions following radiation . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3.9 Active involvement of intestinal stem cells during tissue restitution . . . . . . . . . . . . . . . . . . . . . . . . . 4.3.10 Elevated mesenchymal activities . . . . . . . . . . . . . Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.4.1 Concerns regarding the radiation setup . . . . . . . . . 4.4.2 Impressive regenerative capacity of zebrafish intestine . 4.4.3 Differential sensitivity of intestine to radiation and cancer rate . . . . . . . . . . . . . . . . . . . . . . . . . . 4.4.4 Implications for colorectal cancer therapy . . . . . . . . 4.4.5 Future directions . . . . . . . . . . . . . . . . . . . . . . 124 . 126 . . . . . 130 133 135 135 138 . 139 . 140 . 141 STORM: A General Model to Investigate Stem Cell Number and Their Adaptive Changes 143 5.1 Background . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144 5.2 Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . 146 5.2.1 Development of the STORM model . . . . . . . . . . . . 146 5.2.2 Maintenance of zebrafish . . . . . . . . . . . . . . . . . . 156 5.2.3 Tissue sectioning . . . . . . . . . . . . . . . . . . . . . . 156 5.2.4 Immunohistochemistry . . . . . . . . . . . . . . . . . . . 156 5.3 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158 5.3.1 General characteristics of the crypt-villus system . . . . 158 5.3.2 Determination of the number of epithelial stem cells in a 2D section of the inter-villi pocket of zebrafish (Danio rerio) intestine . . . . . . . . . . . . . . . . . . . . . . . 159 5.3.3 Determination of the stem cell number in each crypt of mouse small intestine . . . . . . . . . . . . . . . . . . . . 163 5.3.4 Determination of the stem cell number in each crypt of human duodenum . . . . . . . . . . . . . . . . . . . . . . 165 5.3.5 Comparison of the intestines of different species . . . . . 168 5.3.6 Uncontrolled expansion of the capacity of stemness upon impaired feedback mechanism . . . . . . . . . . . . . . . 171 5.3.7 Application of the model to help evaluate hyperplasia in human duodenitis and ulcer . . . . . . . . . . . . . . . . 172 5.4 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173 iv 5.4.1 5.5 Epithelium apoptosis is actively initiated in zebrafish intestine before mature cells get exfoliated at the tips of villi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.4.2 Achieving the optimal epithelium renewal rate might be a fundamental principle of the crypt-villus system design by nature . . . . . . . . . . . . . . . . . . . . . . . . . 5.4.3 The number of stem cells is largely conserved in the small intestines of teleost, murine and human . . . . . . . . . 5.4.4 A general model for analysis of stem cell number with equal applicability to teleost, murine and human intestinal tracts . . . . . . . . . . . . . . . . . . . . . . . . . 5.4.5 Homeostasis of intestinal secretory cells takes high priority to ensure the integrity of the feedback mechanism 5.4.6 Growing evidence for validity of the model . . . . . . . Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173 . 174 . 175 . 176 . 176 . 177 . 178 Conclusion 179 6.1 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180 6.2 Future research directions . . . . . . . . . . . . . . . . . . . . . 182 Appendix A 208 v Abstract Unlike the mammalian digestive tract that has been developed into distinct regions for different functions, fish have a relatively simple intestine and many fishes have no recognizable stomach. We used the zebrafish microarray approach to characterize its intestine. By dividing the zebrafish intestine into seven segments along its length, we found that the first five segments resemble the mammalian small intestine and the last two segments resemble the mammalian large intestine. We then investigated the role of Notch signaling and found that a specific group of glycogen-rich fibroblasts were involved in the Notch-mediated cell fate decision process. Further, we studied the effects of radiation and found an interesting pattern of regeneration in the intestine. Moreover, the number of intestinal stem cells was investigated through a novel computational model, which was applicable not only to zebrafish, but also to mammalian intestinal tracts. vi Summary A systemic study of adult zebrafish intestine has been carried out in this thesis project integrating morphological, histological, molecular and computational approaches. Morphologically, the zebrafish intestine is organized as an inverted Z shape in vivo. Dilation of the intestinal tube at its anterior is frequently seen containing ingested food, especially after feeding; while compaction of stools is often observed in its posterior region. Histologically, villi are present almost along the whole intestine with expcetion in the very posterior region (segment S7). Interestingly, crypts are absent along the whole intestine. Our transcriptomic analysis has shown that the zebrafish mucosa only resembles the mouse villi but not the crypts, supporting the absence of crypts in zebrafish. cDNA microarray has been performed to profile the region-specific transcriptomes along the anterior-posterior axis. Transcriptomic analysis shows segments S1-S5 are very similar to each other, while S6 and S7 shows major difference. This is consisten with our qRT-PCR results and in situ hybridization results. Expression of fabp2, the well known marker gene of the small intestine, is switched off in the proximal region of S5. Similar results are seen for villin, another marker gene of the small intestine. vii Dr.24325 Dr.39930 Dr.32463 ZGC:65780 UGT1AB CTSC Zgc:65780 Zgc:123097 Cathepsin C Dr.74638 LOC100007461 Dr.104982 WU:FB55E05 Hypothetical LOC100007461 Wu:fb55e05 Dr.80745 Dr.89609 Dr.74222 Dr.75117 Dr.74031 Dr.82571 Dr.80488 LOC558120 SI:CH211-81I17.1 CKMA SI:DKEY-202B22.2 PPP1R10 Dr.77631 Dr.81898 Dr.24546 protein ZGC:110741 ZGC:66327 Dr.108054 SUPT3H Dr.76556 ZGC:110112 Oogenesis-related protein Hypothetical LOC558120 Si:ch211-81i17.1 Creatine kinase, muscle a Si:dkey-202b22.2 Transcribed locus Protein phosphatase 1, regulatory subunit 10 Zgc:113381 Dr.80679 Dr.86370 Dr.83002 Dr.85461 Dr.119008 Dr.16782 Dr.85829 ZGC:112364 ZGC:77076 ZGC:101657 ZGC:56706 LOC569427 Dr.132294 NOL14 Wu:fa09g06 Transcribed locus Zgc:154063 Dr.110726 Dr.86192 Dr.107641 LOC572168 LOC559127 LOC569277 Dr.80066 Dr.82169 ZGC:123333 Dr.82451 CDC42EP2 CETN2 Dr.91347 ZGC:113381 Dr.32625 Dr.82091 Dr.7724 STKA ZGC:154063 Dr.45587 Dr.33127 ZGC:110611 IFRD2 Dr.79190 ZGC:158607 Zgc:110611 Interferon-related developmental regulator Zgc:158607 Dr.132448 Dr.29131 Transcribed locus Zgc:56476 Dr.143590 Dr.132789 ZGC:85948 ZGC:56476 Dr.105269 SUPT16H Dr.6703 CH211-106H4.12 Dr.85496 FANCL Dr.21244 Dr.82168 UCP2 Dr.84467 ZGC:56388 Suppressor of Ty 16 homolog Fanconi anemia, complementation group L Uncoupling protein Transcribed locus, strongly similar to NP 001004652.1 COX17 cytochrome c oxidase assembly homolog [Danio rerio] Zgc:56388 Dr.79437 Dr.41980 ZGC:110299 ARID3B Dr.80756 Dr.115569 Dr.81807 ZGC:77748 ATXN3 LOC100002531 Dr.9860 MDH1B Dr.75940 DEF Dr.124001 Dr.3552 Dr.122080 Dr.80074 Dr.121869 ZGC:136371 Dr.14064 Dr.51681 ZGC:55418 LOC799717 Dr.78991 LOC100006201 Dr.115855 TK2 POP4 Dr.114689 FAM116B Dr.13689 ZGC:66449 Dr.84866 LOC100005159 Dr.2877 Dr.81115 WU:FB08E07 Dr.42866 MGST1 Transcribed locus Zgc:110741 Hypothetical protein LOC791777 Suppressor of Ty homolog (S. cerevisiae) Hypothetical protein LOC792281 Zgc:112364 Zgc:173915 Zgc:101657 Zgc:56706 Hypothetical LOC569427 Transcribed locus Centrin, EF-hand protein, Hypothetical protein LOC792320 Hypothetical LOC572168 Similar to AWKS9372 Similar to ubiquitously transcribed tetratricopeptide repeat, X chromosome Transcribed locus Zgc:123333 CDC42 effector protein (Rho GTPase binding) Zgc:85948 Transcribed locus, strongly similar to XP 001333536.1 PREDICTED: hypothetical protein [Danio rerio] Hypothetical LOC561742 Hypothetical protein LOC100005159 Wu:fb08e07 Transcribed locus Microsomal glutathione Stransferase Transcribed locus ADP-ribosylation factorlike 4D Zgc:158611 Zgc:64148 Si:dkey-274c14.3 Hypothetical LOC557968 AT rich interactive domain 3B (Bright like) Zgc:77748 Ataxin Hypothetical protein LOC100002531 Malate dehydrogenase 1b, NAD (soluble) Digestive-organ expansion factor Transcribed locus Dr.123167 Dr.76453 ARL4D Dr.7628 Dr.77441 Dr.82147 ZGC:158611 ZGC:64148 SI:DKEY-274C14.3 Dr.78511 LOC559941 Dr.43999 CGNL1 Dr.10250 TDH Zgc:136371 Transcribed locus Hypothetical LOC554559 Transcribed locus, moderately similar to NP 001035095.1 transcription elongation regulator isoform [Homo sapiens] Zgc:55418 Hypothetical protein LOC799717 Hypothetical protein LOC100006201 Hypothetical protein LOC791677 Family with sequence similarity 116, member B Zgc:66449 Dr.133306 Dr.16206 Dr.105356 Dr.77776 CHM ZGC:153334 ZGC:92026 SI:CH211-59D15.5 L-threonine dehydrogenase Choroideremia Zgc:153334 Zgc:92026 Si:ch211-59d15.5 Dr.81544 Dr.116711 LOC561733 Transcribed locus Hypothetical LOC561733 Dr.75239 WU:FB74B10 Wu:fb74b10 Dr.85728 LOC557582 Hypothetical LOC557582 Dr.28581 OPHN1 Dr.121941 Similar to Rho-guanine nucleotide exchange factor Cingulin-like Oligophrenin Transcribed locus 240 Dr.4883 HSD17B4 Dr.18318 SNRPC Dr.36457 PPP2R1B Dr.78320 ZGC:110281 Hydroxysteroid (17-beta) dehydrogenase Small nuclear ribonucleoprotein polypeptide C Protein phosphatase (formerly 2A), regulatory subunit A, beta isoform Zgc:110281 Dr.76567 RGS7 Dr.76198 IM:7150932 Regulator of G-protein signalling Im:7150932 Dr.75142 SI:CH211-197G15.1 Si:ch211-197g15.1 241 Table 8: Genes that are enriched in S7 Unigene Dr.85699 Dr.104972 GeneSym ZGC:158130 Unigene Dr.76238 Dr.79262 Dr.83251 Desc Zgc:158130 Transcribed locus, strongly similar to NP 001001948.1 nucleoporin 54 [Danio rerio] Transcribed locus Dr.122767 Transcribed locus Dr.51681 LOC799717 Hypothetical protein LOC791489 Hypothetical protein LOC792272 Transcribed locus Dr.78991 LOC100006201 Dr.77310 ANXA11A Dr.13660 ZGC:101744 Dr.77138 CPA2 Dr.80069 Dr.105241 SCD Dr.132314 Dr.76148 ZGC:91794 ATP2B1A Dr.83800 DHCR7 Dr.106275 Dr.123334 Dr.133403 Dr.107097 WU:FK81D02 Dr.75792 Dr.123166 Dr.77514 Dr.82353 HOXD9A Dr.77126 Dr.77127 Dr.89216 Dr.115420 Dr.32560 Dr.85873 CTRB1 ZGC:66382 Dr.10050 Dr.134285 Dr.117291 Dr.75974 ADIPOR2 DAO.2 WU:FD10H03 PDZK1L Dr.77685 SLC1A4 Dr.31100 NUDT15 Dr.10201 Dr.79165 Dr.76508 SEPW1 SB:CB14 DIRC2 Dr.1214 ARL6IP1 Dr.21233 Dr.25699 ZGC:103515 ZGC:77082 ELA3L ELA2 TH1L ZGC:113564 EYA4 Dr.81910 Dr.75440 Dr.83156 Dr.78850 Dr.44401 Dr.76387 ZGC:153587 LOC791684 GeneSym SC:D0144 FBXO5 Dr.121869 Desc Sc:d0144 F-box protein Transcribed locus, moderately similar to NP 001035095.1 transcription elongation regulator isoform [Homo sapiens] Hypothetical protein LOC799717 Hypothetical protein LOC100006201 Annexin A11a Dr.76874 TEP1 Dr.78320 ZGC:110281 Telomerase-associated protein Zgc:110281 Dr.80638 Dr.106493 WU:FE05B03 SHCBP1 Wu:fe05b03 Hypothetical LOC554973 Dr.81476 SI:DKEY-171O17.7 Si:dkey-171o17.7 Dr.80835 Dr.76339 SI:CH73-13B6.3 UBE4B Si:ch73-13b6.3 Ubiquitination factor E4B, UFD2 homolog (S. cerevisiae) Hypothetical LOC554619 Serpin peptidase inhibitor, clade B (ovalbumin), member Zgc:92240 Zgc:110687 Tropomyosin (alpha) SWI/SNF related, matrix associated, actin dependent regulator of chromatin, subfamily e, member Transcribed locus Zgc:112971 Zgc:112291 Zgc:153243 Zgc:113294 Zgc:163003 Hypothetical protein LOC792020 Zgc:91794 ATPase, Ca++ transporting, plasma membrane 1a 7-dehydrocholesterol reductase Transcribed locus Transcribed locus Transcribed locus Wu:fk81d02 Dr.47436 Dr.77198 MCM7 SERPINB1 Homeo box D9a Transcribed locus Elastase like Similar to Ela2 protein Dr.84455 Dr.75383 Dr.105413 Dr.75920 ZGC:92240 ZGC:110687 TPM1 SMARCE1 Chymotrypsinogen B1 Zgc:66382 Transcribed locus TH1-like (Drosophila) Zgc:113564 Eyes absent homolog (Drosophila) Adiponectin receptor D-amino-acid oxidase Wu:fd10h03 PDZ domain containing like Solute carrier family (glutamate/neutral amino acid transporter), member Hypothetical protein LOC791620 Selenoprotein W, Sb:cb14 Disrupted in renal carcinoma ADP-ribosylation factorlike interacting protein Zgc:103515 Zgc:77082 Dr.122296 Dr.105858 Dr.45506 Dr.81117 Dr.134857 Dr.13694 ZGC:112971 ZGC:112291 ZGC:153243 ZGC:113294 ZGC:163003 Hypothetical LOC558964 (LOC558964), mRNA Transcribed locus Transcribed locus Transcribed locus Zgc:153587 Hypothetical protein LOC791684 Dr.80584 Dr.15633 Dr.80071 Dr.77534 LOC560382 LOC570432 ZGC:55702 Hypothetical LOC560382 Hypothetical LOC570432 Transcribed locus Zgc:55702 Dr.82867 ZGC:65875 Zgc:65875 Dr.80398 ZGC:153079 Zgc:153079 Dr.87643 Dr.81341 Dr.32320 ZGC:101827 CAPRIN2 LGMN Zgc:101827 Caprin family member Legumain Dr.85513 WU:FC54A11 Wu:fc54a11 Dr.77771 Dr.114174 SI:DKEY-252H13.6 ZGC:63569 Dr.20974 ZGC:55943 Si:dkey-252h13.6 Hypothetical LOC100000446 Zgc:55943 Dr.32415 Dr.6360 Dr.78587 Dr.76985 Dr.80419 WU:FD16G01 TSC1B CLDN10L ZGC:153999 Transcribed locus Wu:fd16g01 Tuberous sclerosis 1b Claudin 10 like Zgc:153999 242 protein Dr.39134 SLC27A1 Dr.6725 Dr.76646 Dr.82756 ZGC:165381 LOC402976 Dr.89589 Dr.78050 Dr.75549 ZGC:101650 ZGC:73324 ZGC:55420 Dr.47389 CEL.2 Dr.84591 Dr.119936 Dr.83470 Dr.107259 ZGC:92392 ZGC:123113 SEPP1A Dr.32109 Dr.77027 ZGC:91959 KNTC2L Dr.31694 ZGC:56304 Dr.79878 DAB2 Dr.77083 Dr.78673 Dr.105771 ZGC:86714 LOC571547 LOC100007704 Dr.78430 Solute carrier family 27 (fatty acid transporter), member Transcribed locus Zgc:165381 Hypothetical protein LOC402976 Zgc:101650 Zgc:73324 Hypothetical protein LOC792156 Hypothetical protein LOC792128 Transcribed locus, strongly similar to NP 001018198.1 spermatogenesis associated 18 [Danio rerio] Zgc:92392 Zgc:123113 Selenoprotein P, plasma, 1a Zgc:91959 Kinetochore associated 2like Zgc:56304 Disabled homolog (Drosophila) Zgc:86714 Hypothetical LOC571547 Similar to Slc7a8-prov protein Transcribed locus, strongly similar to XP 698433.2 PREDICTED: hypothetical protein [Danio rerio] Wu:fj63d08 Transcribed locus Zgc:66317 Transcribed locus, strongly similar to XP 001332873.1 PREDICTED: similar to Solute carrier family 15 (oligopeptide transporter), member [Danio rerio] Similar to LDLR dan Zgc:92139 Zgc:92705 Transcribed locus Nuclear receptor subfamily 5, group A, member TATA box binding protein (Tbp)-associated factor, RNA polymerase I, B Zgc:101841 Dr.76762 IHPK2 Inositol hexaphosphate kinase Dr.81863 Dr.79423 Dr.113263 ZGC:112466 ZGC:114119 LOC793284 Dr.21063 Dr.133000 Dr.75618 NKX3.2 ZGC:171444 Zgc:112466 Zgc:114119 Similar to betamicroseminoprotein NK3 homeobox Transcribed locus Zgc:171444 Dr.78271 ZGC:158414 Zgc:158414 Dr.13175 Transcribed locus Dr.75449 Dr.84876 Dr.74207 LOC555985 CH211-271J4.1 Dr.81839 Dr.91044 ZGC:77563 LOC557719 Dr.1692 ZBTB2B Zinc finger and BTB domain containing 2b Transcribed locus ZGC:113070 Zgc:113070 Transcribed locus Transcribed locus Dr.23725 Dr.83279 Dr.86083 Dr.85095 Dr.121988 Transcribed locus Dr.74013 Dr.23685 Dr.78704 Dr.79959 WU:FJ63D08 Dr.31691 Dr.4044 Dr.85767 Dr.86277 Dr.11921 LOC562438 ZGC:92139 ZGC:92705 Dr.80597 TAF1B Dr.133716 ZGC:101841 Dr.19659 EZRL Ezrin like Dr.80471 ZGC:110537 Dr.78996 ZGC:55661 Hypothetical LOC791913 Zgc:55661 Transcribed locus Dr.76172 FUCA1 Dr.24323 ZGC:56608 Zgc:56608 Dr.117460 LOC792416 Dr.75320 HSPD1 ZGC:162509 WU:FC44A11 Heat shock 60kD protein (chaperonin) Wu:fc44a11 Dr.85731 Dr.76739 Dr.11991 CXXC1 Dr.144128 ZGC:109868 Zgc:109868 Dr.81010 LOC795788 ZGC:66317 NR5A5 Dr.122702 protein Transcribed locus Hypothetical LOC555985 Apoptosis-stimulating protein of p53 Zgc:77563 Hypothetical LOC557719 Dr.79037 Dr.76979 Dr.54293 Dr.105609 ZGC:110259 ZGC:153225 ZGC:110159 Dr.97360 Dr.22874 Dr.12595 Dr.89596 Dr.75252 ZGC:111826 WU:FC21E07 HSF2 ZGC:103559 COL9A2 Dr.77336 ZGC:113196 Dr.10637 LOC797698 Dr.78358 LOC100000526 Dr.76665 LOC566399 Dr.80663 ALG1 Zgc:110259 Zgc:153225 Transcribed locus Zgc:110159 Zgc:111826 Wu:fc21e07 Heat shock factor Zgc:103559 Procollagen, type IX, alpha Zgc:113196 Similar to 5-3 exoribonuclease Hypothetical protein LOC100000526 Hypothetical LOC566399 Asparagine-linked glycosylation homolog (yeast, beta1,4-mannosyltransferase) Fucosidase, alpha-L- 1, tissue Hypothetical protein LOC792416 Zgc:162509 CXXC finger (PHD domain) Similar to CC chemokine SCYA103 243 Dr.78624 Dr.6635 SH3BP5LA Dr.87549 Dr.17300 Dr.80589 Dr.33597 Dr.81772 Dr.79047 Dr.132878 ZGC:112418 LOC559362 SCPEP1 LOC562370 SI:BUSM1-241H12.4 ZGC:113480 LOC100002850 Dr.76472 ZGC:110821 Dr.82787 Dr.117801 IM:7150662 SMAD5 Dr.121617 Dr.81288 MAF Dr.75131 FBL Dr.29762 EFNA2 Dr.75520 GATM Dr.82671 SI:CH211-51L3.4 Dr.85217 Dr.114529 FLJ32675 LOC100003771 Dr.78031 WHSC1 Dr.75811 Dr.107944 INS ABP1 Dr.87101 Dr.84019 Dr.28449 ZGC:110143 ZGC:63614 SLC2A12 Dr.79110 SUFU Dr.76972 ZGC:110109 Dr.9109 Dr.121650 GLMNL Dr.34301 Transcribed locus Dr.24983 SH3-binding domain protein 5-like, a Zgc:112418 Hypothetical LOC559362 Serine carboxypeptidase Hypothetical LOC562370 Si:busm1-241h12.4 Zgc:113480 Similar to Secretory carrier membrane protein 2, like Zgc:110821 Dr.107744 POLR3F Polymerase (RNA) III (DNA directed) polypeptide F Transcribed locus LOC799087 Similar to Oip5 protein Transcribed locus Zgc:73340 Similar to Apoa4 protein Si:dkey-21k10.1 Zgc:101803 Zgc:113183 Dr.117593 Dr.26261 Dr.80630 Dr.132329 Dr.110644 Dr.88420 Dr.45899 ZGC:73340 LOC100004607 SI:DKEY-21K10.1 ZGC:101803 ZGC:113183 Dr.26560 ELOVL5 Dr.76656 Dr.122289 CAPN8 Dr.80332 Dr.3436 MCRS1 LOC100000870 Microspherule protein Hypothetical protein LOC100000870 Dr.14806 PPP2R3C Non-POU domain containing, octamer-binding Glycine amidinotransferase (L-arginine:glycine amidinotransferase) Si:ch211-51l3.4 Dr.6975 ZGC:66107 Protein phosphatase 2, regulatory subunit B”, gamma Zgc:66107 Dr.75913 MAPK3 Mitogen-activated protein kinase Dr.5461 LOC402880 Flj32675 Similar to OTTHUMP00000028706 Wolf-Hirschhorn syndrome candidate protein Preproinsulin Amiloride binding protein (amine oxidase (coppercontaining)) Zgc:110143 Zgc:63614 Solute carrier family (facilitated glucose transporter), member 12 Suppressor of fused homolog (Drosophila) Zgc:110109 Dr.23039 Dr.116914 LOC100005579 Dr.79733 SI:CH211-67E16.9 Hypothetical protein LOC402880 Transcribed locus Hypothetical protein LOC100005579 Si:ch211-67e16.9 Dr.117507 Dr.79151 ZGC:174575 PXK Zgc:174575 PX domain containing serine/threonine kinase Dr.9174 Dr.133159 Dr.37018 ZGC:55673 ZGC:112003 Zgc:55673 Transcribed locus Zgc:112003 Im:7150662 MAD homolog (Drosophila) Transcribed locus V-maf musculoaponeurotic fibrosarcoma (avian) oncogene homolog Fibrillarin ELOVL family member 5, elongation of long chain fatty acids (yeast) Calpain Transcribed locus Dr.75863 ZGC:77488 Zgc:77488 Dr.75558 SLC4A2 Glomulin, like Transcribed locus Dr.122732 Dr.1301 PRKAG1 ITGB5 Integrin, beta Dr.105135 PPRC1 Dr.38076 Dr.81287 ZGC:158780 KRML2 Dr.103297 Dr.123677 LOC564151 Dr.31250 Dr.34264 ZGC:92808 GDPD1 Dr.143610 Dr.81478 WDR43L SULT1ST3 WD repeat domain 43, like Sulfotransferase family 1, cytosolic sulfotransferase Dr.82025 Dr.77381 ZGC:112208 ZGC:92479 Zgc:158780 Kreisler (mouse) mafrelated leucine zipper homolog Zgc:92808 Glycerophosphodiester phosphodiesterase domain containing Zgc:112208 Zgc:92479 Solute carrier family 4, anion exchanger, member Transcribed locus Hypothetical protein LOC791615 Peroxisome proliferatoractivated receptor gamma, coactivator-related Similar to Claspin Transcribed locus Dr.78005 Dr.132401 MPZ SENP3A Dr.84521 Dr.76489 LOC560369 ZGC:92763 Hypothetical LOC560369 Zgc:92763 Dr.132665 Dr.80988 ZGC:153695 PAQR3 Dr.104263 ZGC:92057 Zgc:92057 Dr.6651 PIAS4 Dr.78421 Dr.75949 Dr.4742 ZGC:163008 ZGC:101121 Zgc:163008 Transcribed locus Zgc:101121 Dr.85714 Dr.6944 Dr.133028 ZGC:100860 ADIPOR1A Sc:d0186 SUMO1/sentrin/SMT3 specific peptidase 3a Zgc:153695 Progestin and adipoQ receptor family member III Protein inhibitor of activated STAT, Transcribed locus Zgc:100860 Adiponectin receptor 1a 244 Dr.115882 LOC796920 Hypothetical LOC796920 Dr.81689 Dr.80558 Dr.76364 Dr.107618 Dr.20429 ZGC:55733 ZGC:111879 ZGC:158773 Zgc:55733 Zgc:111879 Zgc:158773 Transcribed locus Discs, large homolog (Drosophila) Im:7136473 DLG7 Dr.109638 ZGC:114129 Dr.31849 Dr.143234 Dr.82465 Dr.5413 ZGC:136380 Dr.84939 ST6GAL1 Dr.132825 ZGC:64095 Dr.114938 H2AFX Dr.114249 HAGH Dr.80461 GNAI2 Dr.39143 Dr.76444 ZGC:112982 ZGC:172180 Dr.78356 Dr.122161 GATA6 Dr.90586 CAMK2G1 Dr.76410 RAB20 Dr.75714 WU:FD20A04 Dr.76827 Dr.79864 LOC556561 Dr.105962 Dr.84470 Dr.143593 Dr.78701 Dr.119225 WU:FC50C06 LOC797345 PCM1 ZGC:110212 LOC797428 ZGC:63528 protein Dr.15041 Dr.80282 Dr.80187 Dr.75118 Dr.78675 Dr.143654 ZGC:110141 ZGC:92035 ZGC:112098 CCNA1 ZGC:77183 Dr.46626 LOC100002480 Zgc:158165 Transcribed locus Zgc:63528 Transcribed locus, moderately similar to XP 520446.2 PREDICTED: similar to ANKRD15 protein [Pan troglodytes] ST6 beta-galactosamide alpha-2,6-sialyltranferase Hypothetical protein LOC792222 H2A histone family, member X Hydroxyacylglutathione hydrolase Guanine nucleotide binding protein (G protein), alpha inhibiting activity polypeptide Zgc:112982 Zgc:172180 Dr.91580 Dr.82327 Dr.77563 Dr.83069 ZGC:171753 ZGC:153958 WU:FB82H05 Dr.75215 ERG Dr.76888 WU:FB37G07 Dr.78720 ZGC:63810 GATA-binding protein Transcribed locus, strongly similar to NP 001104632.1 hypothetical protein LOC559922 [Danio rerio] Calcium/calmodulindependent protein kinase (CaM kinase) II gamma RAB20, member RAS oncogene family Wu:fd20a04 Dr.76645 Transcribed locus, strongly similar to XP 001343059.1 PREDICTED: similar to MGC64297 protein [Danio rerio] Zgc:110141 Zgc:92035 Zgc:112098 Zgc:173602 Zgc:77183 Similar to A kinase (PRKA) anchor protein Zgc:171753 Zgc:153958 Wu:fb82h05 Transcribed locus V-ets erythroblastosis virus E26 oncogene like (avian) Wu:fb37g07 Zgc:63810 Transcribed locus Dr.77740 SETD2 Dr.77595 Dr.83417 BNIP2 Dr.140596 Dr.80631 ZDHHC24 Dr.18206 SET domain containing Transcribed locus BCL2/adenovirus E1B interacting protein Transcribed locus Zinc finger, DHHC-type containing 24 Transcribed locus Dr.5549 ZGC:66286 Zgc:66286 Dr.4573 LGALS9L1 Hypothetical protein LOC791953 Transcribed locus Transcribed locus ZGC:73124 ZGC:100963 ZGC:114123 EPD LOC572175 Zgc:73124 Zgc:100963 Zgc:114123 Ependymin Similar to Muc2 protein Dr.105738 WU:FI38H09 Wu:fi38h09 Dr.79193 MED12 Mediator of RNA polymerase II transcription, subunit 12 homolog Solute carrier family 25 (mitochondrial carrier, phosphate carrier), member 25 Zgc:152928 Globoside alpha-1,3-Nacetylgalactosaminyltransferase 1, like Zgc:63958 Zgc:162290 Zgc:63617 Zgc:56361 Transcribed locus Similar to polyhomeotic like (Drosophila) Wu:fc50c06 Zgc:163126 Pericentriolar material Zgc:110212 Similar to Tubulin folding cofactor B UDP-glucose dehydrogenase Similar to Apoptotic chromatin condensation inducer in the nucleus (Acinus) Far upstream element (FUSE) binding protein Dr.80871 Dr.121558 Dr.106542 SLC25A25 Dr.80632 Dr.77660 Dr.75730 Dr.268 Dr.14396 Dr.113556 UGDH Dr.32211 ACIN1A Dr.76240 FUBP1 Dr.79378 Dr.76512 LOC561658 SEPT7A Hypothetical LOC561658 Septin 7a Dr.84914 Dr.114873 ZGC:152928 GBGT1L3 Dr.79900 Dr.2976 Dr.108624 Dr.78373 CX28.9 SI:CH211-200O3.4 ZGC:86716 Connexin 28.9 Si:ch211-200o3.4 Zgc:86716 Transcribed locus Dr.118849 Dr.16985 Dr.132934 Dr.76745 ZGC:63958 ZGC:162290 ZGC:63617 ZGC:56361 245 Dr.76296 ZGC:56419 Zgc:56419 Dr.82468 TBC1D19 Dr.78711 ACSL4 Dr.78921 Dr.75224 Dr.11428 Dr.80222 ZGC:153420 TBC1 domain family, Dr.120392 member 19 Acyl-CoA synthetase longDr.133630 chain family member Zgc:153420 Dr.13798 Transcribed locus Dr.26907 Similar to supervillin Dr.114009 Cancer susceptibility canDr.78779 didate Similar to NDr.94519 acetylgalactosaminyltransferase Hypothetical protein Dr.83427 LOC791432 Claudin i Dr.75901 Transcribed locus Dr.132230 Glycine dehydrogenase Dr.29995 (decarboxylating) Similar to Chromosome Dr.75243 condensation protein G Hypothetical LOC554697 Dr.96932 Si:dkey-24p1.5 Dr.31566 Transcribed locus Dr.78758 LOC100001855 CASC3 Dr.16810 LOC798400 Dr.9166 BLZF1 Dr.143598 Dr.4955 Dr.76732 GLDC Dr.84109 LOC796103 Dr.23391 Dr.81385 Dr.140471 SLC16A3 SI:DKEY-24P1.5 Dr.4451 Dr.6973 ZGC:55363 WTAP CLDNI Dr.121668 Dr.83415 Dr.79634 LOC561231 Dr.132880 Dr.85634 S100A10A Dr.144133 LOC565165 Dr.123250 Zgc:55363 Wilms tumor associated protein Transcribed locus Transcribed locus Hypothetical LOC561231 Dr.32351 LGALS3L ADA ZGC:110329 DND Transcribed locus Dead end Transcribed locus Transcribed locus ZGC:112296 Zgc:112296 ZGC:91890 Zgc:91890 SI:DKEYP-117H8.4 ZGC:101116 WU:FI27C05 Si:dkeyp-117h8.4 Zgc:101116 Wu:fi27c05 ZGC:113447 Zgc:113447 ZGC:174234 Zgc:174234 Transcribed locus Hypothetical protein LOC792059 Hypothetical LOC554896 Zgc:110788 GPD1 Dr.10580 Dr.83494 NEK2 ZGC:110788 Dr.6833 Dr.25733 Dr.75369 SI:CH211-238N5.5 RDBP DUS1L Transcribed locus S100 calcium binding protein A10a Hypothetical LOC565165 Dr.135199 Dr.87908 IL12A LIFRA Dr.81832 LOC553407 Transcribed locus Dr.76427 Zgc:171298 Zgc:153434 Hypothetical protein LOC798299 Zgc:172295 Cathepsin L.1 Histocompatibility (minor) HA-1 Dr.76152 Dr.78498 Dr.80336 LOC560112 ZGC:77560 CYP11A1 Dr.114385 Dr.77591 Dr.24755 ZGC:163023 SI:DKEY-175G20.1 PPP2R2D Dr.82334 Dr.122543 Dr.18504 NSMCE1 Dr.80441 Dr.14153 Dr.81345 ZGC:171298 ZGC:153434 LOC798299 Dr.90487 Dr.85174 Dr.77065 ZGC:172295 CTSL.1 HMHA1 Dr.47548 Dr.132594 Dr.88756 HOXD11A Dr.72371 WU:FB60G05 Homeo box D11a Transcribed locus Hypothetical protein LOC791572 Wu:fb60g05 Dr.75844 IDH1 Dr.77547 ZGC:100868 Zgc:100868 Dr.84313 ELP3 Dr.36953 ASAH1 Dr.67796 ICN Dr.75608 ESCO2 Dr.24234 SMC2 ZGC:110238 Lectin, galactosidebinding, soluble, (galectin 3)-like Hypothetical protein LOC792200 Zgc:110329 ZGC:153929 Dr.118663 ZGC:64050 Dr.32947 NUPL1 N-acylsphingosine amidohydrolase (acid ceramidase) Hypothetical protein LOC794885 Nucleoporin like Dr.76852 Dr.33734 Dr.55498 Dr.76790 Dr.6471 ZGC:92643 ZGC:110008 WU:FO94F09 RPA1 ZGC:172228 Zgc:92643 Zgc:110008 Wu:fo94f09 Replication protein A1 Zgc:172228 Dr.77600 Dr.42971 Dr.76547 Dr.133660 Dr.16380 SRL ZGC:112455 ZGC:158323 Dr.117314 Dr.111731 Dr.76190 Dr.2636 ZGC:101843 AQP10 CFL1 IMPDH2 Dr.76784 Dr.106681 Dr.79287 Dr.6291 SEPT8A LOC569952 ZGC:110077 Dr.85554 ZGC:112322 Zgc:101843 Aquaporin 10 Cofilin (non-muscle) IMP (inosine monophosphate) dehydrogenase Zgc:112322 Dr.15534 LOC571197 ALDH8A1 Si:ch211-238n5.5 RD RNA binding protein Dihydrouridine synthase 1-like (S. cerevisiae) Interleukin 12a Leukemia inhibitory factor receptor alpha Hypothetical protein LOC100006308 Transcribed locus, moderately similar to NP 071918.1 zinc finger protein 106 homolog [Homo sapiens] Hypothetical LOC560112 Zgc:77560 Cytochrome P450, subfamily XIA, polypeptide Zgc:163023 Si:dkey-175g20.1 Protein phosphatase 2, regulatory subunit B, delta isoform Zgc:153929 Transcribed locus Non-SMC element homolog (S. cerevisiae) Isocitrate dehydrogenase (NADP+), soluble Elongation protein homolog (S. cerevisiae) Ictacalcin Establishment of cohesion homolog (S. cerevisiae) Structural maintenance of chromosomes Sarcalumenin Zgc:112455 Zgc:158323 Transcribed locus Aldehyde dehydrogenase family, member A1 Septin 8a Hypothetical LOC569952 Zgc:110077 Transcribed locus Similar to Thrap4 protein 246 Dr.19939 Dr.9988 Dr.35904 LOC555164 WU:FD44F11 ZGC:85729 Hypothetical LOC555164 Wu:fd44f11 Zgc:85729 Dr.132353 Dr.31086 Dr.91373 RNF14 RNF2 DGAT2 Dr.79955 Dr.2713 ATL3 CD82 Dr.77448 Dr.20969 WDR21 FOXI1 Dr.10893 SARA2 Dr.85037 CHRNE Dr.119277 Dr.78736 Dr.76207 Dr.80393 Dr.122910 ZGC:101026 WU:FC28F08 AQP3 ZGC:66450 Dr.1823 Dr.77932 Dr.8200 Dr.77976 Dr.81932 SI:CH211-150C22.2 SI:CH211-173P18.3 SI:CH211-45M15.2 BYSL GNB3 Dr.84502 Dr.39081 ZGC:103761 ZGC:110239 Atlastin Hypothetical protein LOC792047 Hypothetical protein LOC792159 Zgc:101026 Wu:fc28f08 Aquaporin Zgc:66450 Transcribed locus, moderately similar to NP 956523.1 COP9 signalosome subunit [Danio rerio] Zgc:103761 Zgc:110239 Dr.96184 Dr.11569 ZGC:101095 OSBPL2 Dr.75392 Dr.75265 Dr.117538 Dr.88645 Dr.118834 AK3L1 Dr.79271 Dr.84790 Dr.28227 Dr.7461 Dr.27090 ZGC:77294 UNG IM:7148063 SI:DKEY-114G7.4 DOCK8 Dr.34097 PMP22B Dr.83194 CYP1C1 Dr.2532 PRKCI Adenylate kinase 3-like Transcribed locus Zgc:77785 Zgc:86611 Similar to AHNAK nucleoprotein Peripheral myelin protein 22b Cytochrome P450, family 1, subfamily C, polypeptide Protein kinase C, iota Dr.80082 LOC559441 Dr.114279 Dr.80192 LOC555064 ZGC:162158 Hypothetical LOC555064 Zgc:162158 Dr.122973 Dr.84465 ZGC:110718 Dr.82140 LOC100005813 Dr.76346 Dr.89720 LOC556114 Similar to sno, strawberry notch homolog (Drosophila) Hypothetical LOC556114 Dr.77508 Dr.76643 MAO HIBADHA Dr.79774 CCDC98 Dr.78811 RNASEH2B ZGC:77785 ZGC:86611 LOC566639 Dr.84534 LOC100007999 Dr.3168 SNUPN Dr.116845 Zgc:101095 Oxysterol binding proteinlike Zgc:77294 Uracil-DNA glycosylase Im:7148063 Si:dkey-114g7.4 Similar to dedicator of cytokinesis Hypothetical protein LOC100007999 Snurportin Similar to E3 ubiquitin ligase Transcribed locus Hypothetical protein LOC791932 Transcribed locus Transcribed locus, strongly similar to NP 001073523.1 hypothetical protein LOC571403 [Danio rerio] Importin Hypothetical protein LOC100000528 Transcribed locus Dr.77582 LOC560453 Dr.8058 GBAS Dr.76907 PCBP2 Poly(rC) binding protein Dr.81733 RBBP5 Dr.77012 RCC1 Regulator of chromosome condensation Dr.141683 Dr.107710 LOC561086 Dr.28300 ZGC:66024 Dr.84504 TRAIP Similar to putative adenylate cyclase TRAF-interacting protein Dr.82354 VLDLR Dr.121990 Transcribed locus Dr.79957 TMEM9B Dr.123669 Transcribed locus, strongly similar to NP 001017864.1 hypothetical protein LOC550562 [Danio rerio] RAB30, member RAS oncogene family Lin-7 homolog C (C. elegans) Dr.32244 HSPB1 Dr.78113 ZGC:77390 Zgc:77390 Dr.37073 ZGC:101553 Zgc:101553 RAB30 Dr.52862 LIN7C IPO9 LOC100000528 Cholinergic receptor, nicotinic, epsilon Si:ch211-150c22.2 Si:ch211-173p18.3 Si:ch211-45m15.2 Bystin-like Guanine nucleotide binding protein (G protein), beta polypeptide Monoamine oxidase 3-hydroxyisobutyrate dehydrogenase a Coiled-coil domain containing 98 Ribonuclease H2, subunit B Similar to KIAA1794 Dr.32093 Dr.33564 Dr.133114 Ring finger protein 14 Ring finger protein Diacylglycerol Oacyltransferase WD repeat domain 21 Forkhead box I1 Dr.14824 Dr.84342 ZGC:92085 Zgc:92085 Glioblastoma amplified sequence Retinoblastoma binding protein Transcribed locus, weakly similar to XP 001341760.1 PREDICTED: similar to Si:ch211-14a17.6 [Danio rerio] Zgc:66024 Very low density lipoprotein receptor TMEM9 domain family, member B Heat shock protein, alphacrystallin-related, 247 Dr.1956 Transcribed locus, strongly similar to XP 702446.2 PREDICTED: hypothetical protein [Danio rerio] Zgc:91847 Dr.117581 Adhesion regulating molecule 1b Notch homolog 1a Dr.77391 UTP6 ZGC:154087 Zgc:154087 Dr.86913 FABP6 Dr.82719 DAP3 Fatty acid binding protein 6, ileal (gastrotropin) Death associated protein Dr.32749 ZGC:91847 Dr.104406 ADRM1B Dr.75753 NOTCH1A Dr.31752 HDAC1 Histone deacetylase Dr.76650 ING4 Dr.77619 Dr.1489 Dr.78703 RUVBL1 ZGC:136963 RTKN2 RuvB-like (E. coli) Zgc:136963 Rhotekin Dr.77926 Dr.80524 Dr.119738 ZGC:63774 WU:FI34B01 SETB Dr.132305 Dr.80724 ZGC:77439 SULT1ST6 Dr.80032 Dr.104887 ZGC:162976 LOC572121 Dr.122409 Dr.17174 Dr.31639 ANKRD28 ZGC:112095 Zgc:77439 Sulfotransferase family, cytosolic sulfotransferase Transcribed locus Ankyrin repeat domain 28 Zgc:112095 Dr.134550 Dr.78888 Dr.80713 ZGC:77906 ZGC:77415 FSHR Dr.77202 RDH1L Retinol dehydrogenase 1, like Dr.106217 C20ORF20 Dr.106380 PRDM8 PR domain containing Dr.86476 LOC100001302 Dr.116756 RIPK4 Receptor-interacting serine-threonine kinase Dr.140573 Transcribed locus Dr.104286 Dr.37970 Dr.79014 SI:RP71-1C10.3 Dr.115188 Dr.82751 SERINC5 SSX2IP Dr.77343 Dr.75547 Dr.140810 TRY Dr.77961 ZGC:92765 Trypsin Transcribed locus, weakly similar to XP 690938.2 PREDICTED: hypothetical protein [Danio rerio] Transcribed locus Cytochrome P450, family 2, subfamily J, polypeptide 30 Choline phosphotransferase Acyl-CoA synthetase longchain family member Zgc:92765 Dr.132611 LOC793280 Dr.40298 Dr.77610 Dr.80572 WU:FB58E08 ZGC:101667 ZGC:103530 Wu:fb58e08 Zgc:101667 Zgc:103530 Dr.120800 Dr.41494 Dr.74211 ZGC:92512 WU:FI46G11 TAKRP Dr.37643 Dr.2274 ZGC:103509 STARD3 Zgc:103509 START domain containing Zgc:55292 Dr.114283 Dr.21958 WU:FD16E03 LBR Dr.78401 AATF Golgi autoantigen, golgin subfamily a, Hepatoma-derived growth factor, related protein Dr.77471 Dr.105427 Dr.37032 CYP2J30 Dr.75843 CHPT1 Dr.79066 ACSL1 Dr.106021 SLC43A1 Dr.85700 NUP43 Dr.1786 ZGC:55292 Dr.79972 GOLGA5 Dr.75202 HDGFRP2 Dr.75603 H3F3A H3 histone, family 3A Dr.83735 HPRT1L Dr.76761 Dr.2143 ZGC:91996 PRPF38B Zgc:91996 PRP38 pre-mRNA processing factor 38 (yeast) domain containing B Dr.31063 Dr.132378 ZGC:86895 PLS3 Dr.93960 UTP6, small subunit (SSU) processome component, homolog (yeast) Inhibitor of growth family, member Zgc:109744 Wu:fi34b01 SET translocation (myeloid leukemiaassociated) B Zgc:162976 Similar to XL-INCENP Zgc:77906 Zgc:77415 Follicle stimulating hormone receptor Chromosome 20 open reading frame 20 (H. sapiens) Similar to ubiquitinactivating enzyme E1 Transcribed locus, weakly similar to NP 081830.2 ubiquitin specific peptidase 38 [Mus musculus] Transcribed locus, strongly similar to NP 001071010.2 mitochondrial methionyl-tRNA formyltransferase [Danio rerio] Si:rp71-1c10.3 Transcribed locus Serine incorporator Synovial sarcoma, X breakpoint interacting protein Solute carrier family 43, member Nucleoporin 43 Hypothetical protein LOC793280 Zgc:92512 Wu:fi46g11 T-cell activation kelch repeat protein Wu:fd16e03 Lamin B receptor Apoptosis antagonizing transcription factor Transcribed locus Transcribed locus, strongly similar to NP 998703.1 v-crk sarcoma virus CT10 oncogene-like [Danio rerio] Hypoxanthine phosphoribosyltransferase 1, like Zgc:86895 Plastin (T isoform) 248 Dr.132490 ZGC:77867 Zgc:77867 Dr.79720 Dr.133457 ZGC:86839 Zgc:86839 CDNA clone AGE:7053246 Dr.24235 DDX18 Dr.7003 BMP15 Dr.116102 Dr.77703 ELA2L UHRF1 Dr.84043 Dr.5696 ZGC:86635 PRC1 Dr.32573 Dr.21094 Transcribed locus, weakly similar to XP 001333287.1 PREDICTED: similar to ENSANGP00000022061 isoform [Danio rerio] Transcribed locus Similar to transcription factor-like nuclear regulator Phosphoribosyl pyrophosphate synthetaseassociated protein Serine/threonine kinase 38 like Nucleolar protein 10 Similar to Protein phosphatase 1, regulatory (inhibitor) subunit 3B Wu:fd14a01 ATPase, Na+/K+ transporting, alpha 3b polypeptide Transcribed locus, moderately similar to NP 001107527.1 hypothetical protein LOC100135392 [Xenopus tropicalis] Heterogeneous nuclear ribonucleoprotein L Zgc:92689 Dr.81955 Dr.81616 LOC569587 DEAD (Asp-Glu-Ala-Asp) box polypeptide 18 Dr.106367 PRPSAP2 Bone morphogenetic protein 15 Elastase like Ubiquitin-like, containing PHD and RING finger domains, Zgc:86635 Protein regulator of cytokinesis Dr.78235 STK38L Dr.4705 Dr.119787 NOL10 LOC554474 Dr.78602 Dr.122142 WU:FD14A01 ATP1A3B RPL11 Ribosomal protein L11 Dr.135679 Dr.82263 LOC796453 HNRNPL POLA1 Hypothetical protein LOC796453 Polymerase (DNA directed), alpha Transcribed locus, strongly similar to NP 956291.1 TAR (HIV) RNA binding protein [Danio rerio] Transcribed locus, strongly similar to XP 001332593.1 PREDICTED: similar to L(3)mbt-like (Drosophila) isoform [Danio rerio] MYST histone acetyltransferase Hypothetical protein LOC799140 Transcribed locus, strongly similar to XP 001341031.1 PREDICTED: hypothetical protein [Danio rerio] Zgc:100869 Dr.118088 Dr.75324 Dr.82744 ZGC:92689 Dr.80840 LOC564852 Similar to phosphofructo-2kinase/fructose-2,6biphosphatase Dr.81780 PANE1 Proliferation associated nuclear element Dr.120681 ZGC:171428 Zgc:171428 Dr.106432 ZGC:171537 Zgc:171537 Dr.14778 ZGC:103752 Zgc:103752 Dr.76520 ST13 Forkhead box C1b Chromosome open reading frame 24 Dr.80701 Dr.12565 VAT1 Suppression of tumorigenicity 13 (colon carcinoma) (Hsp70 interacting protein) Transcribed locus Vesicle amine transport protein homolog (T californica) Hypothetical protein LOC100004500 Aryl hydrocarbon receptor nuclear translocatorlike 1a Transcribed locus Integrator complex subunit Transcribed locus Sperm associated antigen Hypothetical protein LOC100005281 Cytoskeleton associated protein Zgc:110200 Hypothetical LOC561668 Dr.140642 Dr.142263 Dr.77649 MYST2 Dr.106465 LOC799140 Dr.3839 IM- Dr.75741 ZGC:100869 Dr.83301 Dr.75260 FOXC1B C2ORF24 Dr.85085 PDLIM7 PDZ and LIM domain Dr.121512 LOC100004500 Dr.80637 ITGA5 Dr.75458 ARNTL1A Dr.81184 Dr.82521 ZGC:92818 ZGC:73144 Dr.78902 Dr.82999 INTS6 Dr.80659 Dr.75229 WU:FI33G05 Integrin, alpha (fibronectin receptor, alpha polypeptide) Zgc:92818 Hypothetical protein LOC792240 Wu:fi33g05 Transcribed locus Dr.132373 Dr.80821 SPAG6 Dr.77760 PUS1 Dr.17457 ZGC:63667 Dr.88329 HIF1AL2 Dr.75286 CKAP5 Dr.75372 Dr.75748 STARD3NL ORC6L Dr.116678 Dr.67263 ZGC:110200 LOC561668 Hypothetical protein LOC792133 Hypoxia-inducible factor 1, alpha subunit, like STARD3 N-terminal like Origin recognition complex, subunit homologlike (yeast) 249 6- Dr.80266 ITM2C Dr.85627 ANXA3A Dr.82056 LHX8 LIM homeobox LOC567390 Integral membrane protein 2C Hypothetical protein LOC791593 Hypothetical LOC567390 Dr.118555 Dr.79519 ATP5S Dr.114001 ZGC:110183 Zgc:110183 Dr.119419 LOC572149 Dr.79757 Dr.81511 Dr.76676 ZGC:77112 ZGC:113156 LOC793260 Zgc:77112 Zgc:113156 Hypothetical protein LOC793260 Hypothetical protein LOC100001846 Inhibitor of growth family, member 5b Dr.13909 Dr.59 Dr.75602 ANXA1A TRIM71 Dr.42600 LOC553397 Dr.20155 SS18 Transcribed locus Dr.82585 LOC562304 Poly(A) polymerase gamma Rac GTPase-activating protein Dr.79823 ZGC:91926 ATP synthase, H+ transporting, mitochondrial F0 complex, subunit s Similar to Dihydropyrimidine dehydrogenase Transcribed locus Annexin A1a Tripartite motifcontaining 71 Hypothetical protein LOC553397 Synovial sarcoma translocation, chromosome 18 (H. sapiens) Similar to cytochrome P450, family 2, subfamily J, polypeptide Zgc:91926 Dr.16483 SLC39A6 Zgc:56412 Similar to cell surface flocculin Anillin, actin binding protein-like Hypothetical LOC569148 Cyclin-dependent kinase Dr.15227 Dr.80564 RBM38 Dr.114476 ZGC:92161 Dr.80447 Dr.75906 ZGC:55983 SLC25A3 Dr.133005 LOC100001846 Dr.79544 ING5B Dr.75255 Dr.79169 PAPOLG Dr.105341 RACGAP1 Dr.23593 Dr.106684 ZGC:56412 LOC799913 Dr.74227 ANLN-LIKE Dr.40624 Dr.75152 LOC569148 CDK2 Dr.7036 G3BP1 Dr.106515 LOC795458 Dr.79840 Dr.79258 ZGC:92006 LOC100000846 Dr.67738 Dr.41866 Dr.15775 ZGC:153795 CRYBB3 DKEY-151P17.3 Dr.75966 Dr.83578 THOC1 LOC556669 Dr.52170 Dr.78694 UTP15 Dr.13960 S100A1 Dr.118684 LOC100002310 Dr.77945 DNMT5 GTPase activating protein (SH3 domain) binding protein Similar to ENSANGP00000022061 Zgc:92006 Similar to PHD finger protein 12 Zgc:153795 Crystallin, beta B3 Plasma membrane proteolipid THO complex Hypothetical LOC556669 Dr.77751 Transcribed locus Dr.121634 Dr.79728 LOC564287 Dr.106024 Dr.17275 WU:FC46H12 WU:FI75F03 Dr.80157 Dr.79711 Dr.82376 ZGC:56161 ZGC:158802 ZGC:63863 Dr.99488 Dr.39528 ZGC:171912 LOC100001396 Transcribed locus, strongly similar to NP 001002726.1 WD repeat and HMG-box DNA binding protein [Danio rerio] Utp15, U3 small nucleolar ribonucleoprotein, homolog S100 calcium binding protein A1 Hypothetical protein LOC100002310 Dr.7489 DNA (cytosine-5-)methyltransferase Dr.74706 Dr.2195 Similar to MGC69156 protein Wu:fc46h12 Wu:fi75f03 Zgc:56161 Zgc:158802 Hypothetical protein LOC797468 Zgc:171912 Hypothetical protein LOC100001396 Transcribed locus, weakly similar to NP 001034277.1 ras responsive element binding protein isoform [Mus musculus] TNPO3 Transportin Dr.120175 HSD11B3 Dr.82458 DDX26B Hydroxysteroid (11-beta) dehydrogenase DEAD/H (Asp-Glu-AlaAsp/His) box polypeptide 26B Transcribed locus, moderately similar to NP 775564.1 solute carrier family 43, member [Mus musculus] Hypothetical protein LOC794025 WW domain binding protein Si:ch211-51n14.2 Dr.109592 ZGC:92151 Zgc:92151 Dr.116738 LOC794025 Dr.86190 ZGC:86715 Zgc:86715 Dr.8749 WBP2 Dr.80969 LOC100004989 SI:CH211-51N14.2 ZGC:100856 Hypothetical LOC100004989 Zgc:100856 Dr.117431 Dr.80848 Dr.86027 LOC100006639 protein Solute carrier family 39 (zinc transporter), member Transcribed locus RNA binding motif protein 38 Hypothetical protein LOC791539 Zgc:55983 Solute carrier family 25 (mitochondrial carrier, phosphate carrier), member Transcribed locus Hypothetical LOC100006639 250 protein Dr.77864 PHC2 Dr.82476 Dr.33507 ZGC:112072 ZGC:123170 Polyhomeotic-like (Drosophila) Zgc:112072 Zgc:123170 Dr.67664 SI:DKEY-8L13.4 Si:dkey-8l13.4 Dr.78239 DDI2 Dr.47041 ZGC:92126 DNA-damage protein Zgc:92126 Dr.67618 USP25 Dr.29859 Dr.80409 IM:7142837 IVNS1ABPB Dr.394 inducible Ubiquitin specific protease 25 Im:7142837 Influenza virus NS1A binding protein b Transcribed locus, strongly similar to XP 700078.2 PREDICTED: similar to D13S106E-like [Danio rerio] Parvalbumin Dr.2885 FTSJ1 FtsJ homolog (E. coli) Dr.76280 Dr.87817 ZGC:152779 DNAJC5 Dr.25529 PHEX Dr.132507 ZGC:85696 Zgc:152779 DnaJ (Hsp40) homolog, subfamily C, member Phosphate regulating gene with homologues to endopeptidases on the X chromosome Zgc:85696 Dr.75477 RTF1 Dr.29744 SP8L Dr.40661 Dr.8916 ZGC:110734 Rtf1, Paf1/RNA polymerase II complex component, homolog (S. cerevisiae) Sp8 transcription factorlike Transcribed locus Zgc:110734 Dr.32297 CAHZ Carbonic anhydrase Dr.78166 PVALB7 Dr.81229 Dr.79083 Dr.4660 TMEM57 LOC100006592 Transmembrane protein 57 Hypothetical protein LOC100006592 Dr.132872 Dr.9441 Dr.32436 ZGC:101883 Dr.89166 Dr.32150 Dr.119791 DJ383J4.3L Dr.75610 BHMT ANKRD12 UTXL1 Dr.86041 ZGC:92277 Hypothetical protein LOC792171 DJ383J4.3-like Transcribed locus, strongly similar to XP 001332437.1 PREDICTED: hypothetical protein [Danio rerio] Betaine-homocysteine methyltransferase Zgc:92277 Dr.77969 LOC796797 Dr.75815 FOXA Forkhead box A sequence Dr.50820 LOC565671 Dr.78299 Dr.80537 Dr.30247 PHF17 ZGC:55621 ZGC:77118 PHD finger protein 17 Zgc:55621 Zgc:113969 Dr.89787 Dr.123739 Dr.80660 ZGC:113334 Dr.86116 ZGC:56589 Dr.83896 LOC100005060 Dr.81259 Dr.82958 DAZL MCEE Dr.121806 Dr.113808 PLS1 Dr.76154 ZGC:77241 Novel protein similar to vertebrate phosphatidylinositol glycan anchor biosynthesis, class A (paroxysmal nocturnal hemoglobinuria) (PIGA, zgc:56589) Daz-like gene Methylmalonyl CoA epimerase Zgc:77241 Dr.77001 CDC42EP4 Dr.87644 ZGC:113259 Zgc:113259 Dr.82697 Dr.84074 Dr.36081 Dr.78997 Dr.75779 Dr.15518 SB:CB157 KIF11 Sb:cb157 Kinesin family member 11 Transcribed locus Pl10 Zgc:113343 Dr.115947 Dr.80478 Dr.79117 Dr.37370 Dr.4322 PL10 ZGC:113343 Transcribed locus, strongly similar to XP 001332444.1 PREDICTED: similar to sreb2 isoform [Danio rerio] Ankyrin repeat domain 12 Ubiquitously transcribed tetratricopeptide repeat, X chromosome like Transcribed locus Dr.80967 Dr.84741 ZGC:112254 ZGC:113085 Zgc:112254 Zgc:113085 Dr.114161 ZGC:112397 Zgc:112397 Hypothetical protein LOC796797 Similar to MGC89155 protein Zgc:113334 Transcribed locus Polo-like kinase (Drosophila) Hypothetical protein LOC100005060 PLK4 LOC563252 ZGC:55308 SI:DKEY-97O5.1 ZGC:103562 DKEYP-94H10.2 Transcribed locus Plastin (I isoform) CDC42 effector protein (Rho GTPase binding) Transcribed locus, moderately similar to XP 001345745.1 PREDICTED: hypothetical protein [Danio rerio] Similar to MTG16a Zgc:55308 Si:dkey-97o5.1 Zgc:103562 Novel protein similar to vertebrate PAS domain containing serine/threonine kinase (PASK) 251 Dr.114177 DNAJA3A Dr.47266 SNIP1 Dr.105744 SI:CH211-152C12.2 DnaJ (Hsp40) homolog, subfamily A, member 3A Smad nuclear interacting protein Si:ch211-152c12.2 Dr.52195 ZGC:175186 Dr.82434 SLA/LPL Dr.77921 LOC564669 Similar to ARVCF Dr.77353 ZGC:101663 Dr.83148 LOC556113 LOC558028 ZGC:63504 Similar to FLJ00281 protein Zgc:63504 Dr.116962 Dr.76348 Dr.133285 ZGC:153878 Dr.143375 Zgc:175186 Transcribed locus Soluble liver antigen/liver pancreas antigen (Homo sapiens), like Hypothetical protein LOC792193 Similar to exosome component 10 Hypothetical protein LOC795094 Embryonic ectoderm development Zgc:65845 Dr.104975 ZGC:56513 Zgc:56513 Dr.83484 EED Dr.18312 U2AF1 Dr.120811 ZGC:65845 Dr.6442 Dr.82342 Dr.80456 ZGC:92875 DKEY-57A22.11 Dr.36307 Dr.14549 Dr.25277 LOC562529 ZGC:92307 AGR2 Hypothetical LOC562529 Zgc:92307 Anterior gradient homolog (Xenopus laevis) Dr.76462 RBB4 U2(RNU2) small nuclear RNA auxiliary factor Zgc:92875 Similar to CG14692-PA Transcribed locus, strongly similar to XP 691447.2 PREDICTED: hypothetical protein [Danio rerio] Retinoblastoma binding protein Transcribed locus Zgc:110586 Dr.107002 Dr.106921 ZGC:110586 Dr.35596 Dr.115906 TPP1 LOC799556 Dr.75192 Dr.120542 ZGC:109901 ZGC:63920 Dr.83536 ZGC:103540 Zgc:103540 Dr.120243 Dr.114355 SI:CH211-197I12.2 Dr.78105 Dr.15263 ZGC:56518 ZGC:73231 Si:ch211-197i12.2 Transcribed locus, strongly similar to NP 999931.1 mediator of RNA polymerase II transcription, subunit 25 [Danio rerio] Zgc:56518 Zgc:73231 Dr.140985 Dr.16383 ZGC:112365 LOC564722 Similar to LOC559853 protein Bridging integrator Dr.79098 MBD1 Dr.6354 CNOT3 Tripeptidyl peptidase I Similar to LOC553285 protein Zgc:109901 Zgc:63920 Dr.113957 LOC100007887 Dr.37659 BIN2 Dr.70549 ARL8 ADP-ribosylation like factor- Dr.143602 Dr.4206 ZGC:76977 Dr.133834 Dr.29879 Hypothetical protein LOC792029 Transcribed locus Dr.78519 SPAG1 Dr.77026 Dr.11520 SI:CH211-51E12.7 Transcribed locus Si:ch211-51e12.7 Dr.77928 Dr.6680 SCYL3 MKNK2A Dr.75704 PTGDS Prostaglandin D2 synthase Dr.76983 ZGC:110286 Dr.83502 ZGC:56310 Zgc:56310 Dr.80737 ZGC:110307 Transcribed locus Dr.43244 Dr.77272 Dr.24982 E2F4 ZGC:56585 Dr.132634 HERPUD1 Dr.23244 ZGC:66359 Dr.132340 Dr.77971 LOC798926 Dr.78423 LOC571567 Transcribed locus Hypothetical protein LOC798926 Hypothetical LOC571567 Dr.77484 ZGC:110417 Zgc:110417 Dr.76573 SPNS1 Dr.75662 ZGC:114087 Zgc:114087 Dr.76353 TIAL1 Dr.77172 ZGC:153968 Zgc:153968 Dr.5024 PHF16 Zgc:112365 Similar to receptor associated protein 80 Methyl-CpG binding domain protein CCR4-NOT transcription complex, subunit Transcribed locus, strongly similar to XP 001336617.1 PREDICTED: similar to Adenylate kinase 3-like [Danio rerio] Zgc:66359 Sperm associated antigen SCY1-like (S. cerevisiae) MAP kinase-interacting serine/threonine kinase 2a Hypothetical protein LOC791833 Hypothetical protein LOC791571 Transcribed locus, strongly similar to NP 001017717.1 gammabutyrobetaine hydroxylase [Danio rerio] E2F transcription factor Hypothetical protein LOC792146 Homocysteine-inducible, endoplasmic reticulum stress-inducible, ubiquitin-like domain member Spinster homolog (Drosophila) TIA1 cytotoxic granuleassociated RNA binding protein-like PHD finger protein 16 252 Dr.83525 ZGC:162268 Zgc:162268 Dr.78256 IVNS1ABPA Dr.105855 Dr.77985 ZGC:171818 ZGC:92279 Influenza virus binding protein a Zgc:171818 Zgc:92279 Dr.14734 PUS7 Dr.90256 DSCC1 Dr.80201 Dr.76675 ALDH3D1 Zgc:64130 Zgc:103747 Dr.47567 Dr.132384 SI:DKEY-30H14.2 SULT1ST1 Dr.81681 Dr.122970 Dr.77434 ZGC:55557 ZGC:56326 Dr.32820 Dr.75970 ZGC:56597 FLNA Zgc:56597 Filamin A, alpha Dr.85904 Dr.39952 IM:7143992 VPS33A BCL7A ZGC:66488 Sorting nexin 33 Transcribed locus Similar to zona pellucida glycoprotein ZPB Zgc:171776 Hypothetical protein LOC791435 Zgc:55879 Rho guanine nucleotide exchange factor (GEF) 7b B-cell CLL/lymphoma 7A Hypothetical LOC555138 Transcribed locus Aldehyde dehydrogenase family, member D1 Si:dkey-30h14.2 Sulfotransferase family, cytosolic sulfotransferase Zgc:55557 Transcribed locus Zgc:56326 Dr.78765 Dr.36931 ZGC:64130 ZGC:103747 Dr.90996 Dr.133388 Dr.78823 SNX33 Dr.116371 Dr.75241 ZGC:171776 PCID2 Dr.116223 Dr.78916 ZGC:55879 ARHGEF7B Dr.79340 Dr.80627 ZGC:56653 B3GAT3 Dr.133796 Dr.30339 ZGC:162319 ZGC:85911 Zgc:162319 Zgc:85911 Dr.35822 Dr.83245 ZGC:92425 SKP1 Dr.77853 SRPK1 Dr.80551 ZGC:110655 Dr.104630 MBD3A Dr.83159 IM:7140357 Im:7140357 Dr.88388 NEDD8L Dr.256 SEC61B SEC61, beta subunit Dr.117303 Dr.109645 Dr.80703 CRYGMX ZGC:152986 ATXN7L2 Serine/arginine-rich protein specific kinase Methyl-CpG binding domain protein 3a Neural precursor cell expressed, developmentally down-regulated 8, like Crystallin, gamma MX Zgc:152986 Ataxin 7-like Im:7143992 Vacuolar protein sorting 33A Zgc:56653 Beta3glucuronyltransferase Zgc:92425 S-phase kinase-associated protein Zgc:110655 Dr.28229 Dr.78604 Dr.75185 Dr.36499 Dr.75231 ZGC:110767 ZGC:101635 CHERP Dr.81104 LOC568795 RCL1 ZGC:92027 ZGC:113828 Dr.87304 Dr.81961 ZGC:77234 LOC100002864 Dr.78406 Dr.117779 ZGC:66448 ZGC:110128 Dr.72357 Dr.117029 ZGC:153136 ZGC:112265 Dr.75159 Dr.11480 WU:FB25B09 RAP2IP Similar to Anaphase promoting complex subunit Zgc:92027 Hypothetical protein LOC797940 Zgc:66448 Hypothetical protein LOC791907 Wu:fb25b09 Rap2 interacting protein Dr.31536 Dr.79516 Dr.106451 Zgc:110767 Zgc:101635 Calcium homeostasis endoplasmic reticulum protein RNA terminal phosphate cyclase-like Zgc:77234 Hypothetical protein LOC100002864 Zgc:153136 Zgc:112265 Dr.122414 Dr.118526 LOC556210 Dr.78825 ELOVL1 Dr.96078 ZGC:100836 Dr.13043 TPST1L Dr.140317 Dr.83943 Dr.79979 ZGC:110333 ZGC:113100 Hypothetical protein LOC791805 Tyrosylprotein sulfotransferase 1, like Zgc:110333 Zgc:113100 Transcribed locus Dr.77086 Dr.105425 Dr.12491 ZGC:92664 ZGC:92164 LOC562579 Transcribed locus Dr.77870 RNGTT Dr.118097 Dr.143376 LOC566888 ZGC:86753 Zgc:92664 Zgc:92164 Similar to complement C42 RNA guanylyltransferase and 5’-phosphatase Hypothetical LOC566888 Zgc:86753 Dr.115125 ZGC:77336 Zgc:77336 Dr.77116 ZGC:101540 Zgc:101540 Dr.45962 Dr.45332 Dr.82411 SPTLC2 Dr.72465 ZGC:153103 LOC572451 Dr.75346 Dr.123514 Dr.80056 NS1A Dr.119113 SUV420H1 Dr.114210 LOC794370 Dr.85121 Dr.122845 Dr.83945 ZGC:56259 BXDC5 Transcribed locus Dehydrodolichyl diphosphate synthase CDNA clone IMAGE:7897837 Suppressor of variegation 4-20 homolog (Drosophila) Similar to DNA polymerase delta1 catalytic subunit Zgc:56259 Transcribed locus Brix domain containing Dr.78482 IM:7158730 Im:7158730 DHDDS Dr.51813 Dr.122815 Pseudouridylate synthase homolog (S. cerevisiae) Zgc:103507 Transcribed locus S100 calcium binding protein V1 Zgc:100836 Transcribed locus Dr.86126 Transcribed locus LOC792966 Similar to cathepsin A Transcribed locus Serine palmitoyltransferase, long chain base subunit Zgc:153103 253 Dr.121695 Dr.123021 Dr.140590 Dr.22945 LOC795535 Dr.31447 MBTPS2 Dr.29035 CTNNB2 Dr.7577 Dr.77837 LOC100005775 Dr.20277 ACTA2 Dr.133827 Dr.24325 ZGC:63676 ZGC:65780 Dr.32463 Dr.115301 CTSC LOC567840 Dr.80745 Dr.143767 Dr.14063 ZGC:152922 EML2 Dr.74222 SI:CH211-81I17.1 Dr.82571 Transcribed locus, moderately similar to NP 001117721.1 glucokinase [Oncorhynchus mykiss] Transcribed locus CDNA clone MGC:173911 IMAGE:5915517 Hypothetical protein LOC795535 Membrane-bound transcription factor protease, site Catenin, beta Dr.91026 LOC100006536 Hypothetical LOC100006536 protein Dr.80532 Dr.85930 DCP1A ZGC:66419 Dr.104703 U2AF2A Dr.38382 NBEA Decapping enzyme Hypothetical protein LOC791547 U2 small nuclear RNA auxiliary factor 2a Neurobeachin Dr.85608 TBPL2 Transcribed locus, strongly similar to XP 691993.2 PREDICTED: similar to pogo transposable element with ZNF domain [Danio rerio] Similar to MGC53357 protein Actin, alpha 2, smooth muscle, aorta Zgc:63676 Zgc:65780 Dr.81537 ZGC:101021 TATA box binding protein like Zgc:101021 Dr.81898 ZGC:110741 Zgc:110741 Dr.24546 ZGC:66327 Dr.87085 Dr.108054 ZGC:92113 SUPT3H Cathepsin C Similar to KIAA1573 protein Oogenesis-related protein Zgc:152922 Echinoderm microtubule associated protein like Si:ch211-81i17.1 Dr.80679 Dr.84245 ZGC:112364 CUL4A Hypothetical protein LOC791777 Zgc:92113 Suppressor of Ty homolog (S. cerevisiae) Zgc:112364 Cullin 4A Dr.31762 Dr.86370 Dr.36480 WU:FI12A09 ZGC:77076 ZGC:103645 Wu:fi12a09 Zgc:173915 Zgc:103645 Dr.77910 FAM46C Transcribed locus Dr.31094 RBKS Dr.119008 LOC569427 Family with sequence similarity 46, member C Hypothetical protein LOC792027 Hypothetical LOC569427 Dr.531 DCPS MRNA decapping enzyme Dr.16782 Dr.85829 CETN2 Dr.76341 LOC100002247 Dr.80488 PPP1R10 Dr.75902 Dr.91347 ZGC:158387 ZGC:113381 Hypothetical protein LOC100002247 Protein phosphatase 1, regulatory subunit 10 Zgc:158387 Zgc:113381 Dr.32625 STKA Wu:fa09g06 Dr.132294 NOL14 Dr.82091 Dr.45587 Dr.85340 ZGC:110611 MRPL14 Dr.13677 Dr.110726 Dr.86192 ZGC:55317 LOC572168 LOC559127 Dr.33127 IFRD2 Transcribed locus Zgc:110611 Mitochondrial ribosomal protein L14 Interferon-related developmental regulator Dr.107641 LOC569277 Dr.83423 LOC569187 Dr.82169 ZGC:123333 Dr.1220 ZGC:64155 Similar to PHD finger protein Zgc:64155 Dr.82451 CDC42EP2 Transcribed locus Myosin regulatory light chain interacting protein Fanconi anemia, complementation group L Uncoupling protein Dr.143590 Dr.84866 ZGC:85948 LOC100005159 Dr.76453 ARL4D Transcribed locus, strongly similar to NP 001004652.1 COX17 cytochrome c oxidase assembly homolog [Danio rerio] Hypothetical LOC557968 Protein arginine Nmethyltransferase AT rich interactive domain 3B (Bright like) Zgc:77748 Dr.119324 ZGC:110245 ADP-ribosylation like 4D Zgc:110245 Dr.7628 Dr.77441 ZGC:158611 ZGC:64148 Zgc:158611 Zgc:64148 Dr.82147 SI:DKEY-274C14.3 Si:dkey-274c14.3 Dr.78511 LOC559941 Similar to Rho-guanine nucleotide exchange factor Dr.122884 Dr.75665 MYLIP Dr.85496 FANCL Dr.21244 UCP2 Dr.82168 Dr.79437 Dr.3855 ZGC:110299 PRMT7 Dr.41980 ARID3B Dr.80756 ZGC:77748 Dr.123167 Transcribed locus Centrin, EF-hand protein, Hypothetical protein LOC792320 Zgc:55317 Hypothetical LOC572168 Similar to AWKS9372 Similar to ubiquitously transcribed tetratricopeptide repeat, X chromosome Zgc:123333 CDC42 effector protein (Rho GTPase binding) Zgc:85948 Hypothetical protein LOC100005159 Transcribed locus 254 factor- Dr.27180 PPARB2 Dr.86792 DT1P1A10L Dr.115569 Dr.81807 ATXN3 LOC100002531 Dr.53929 Dr.75940 SNF1LK2B DEF Dr.124001 Dr.122080 Dr.80074 Dr.132767 Dr.33631 POP4 ZGC:77051 SI:CH211-124K10.2 Peroxisome proliferator activated receptor beta Hypothetical protein DT1P1A10 (human) - like Ataxin Hypothetical protein LOC100002531 SNF1-like kinase 2b Digestive-organ expansion factor Transcribed locus Transcribed locus Hypothetical LOC554559 Zgc:77051 Si:ch211-124k10.2 Dr.43999 CGNL1 Cingulin-like Dr.84960 GRTP1B Dr.133306 Dr.81558 CHM SI:DKEY-98F17.2 Hypothetical LOC791767 Choroideremia Si:dkey-98f17.2 Dr.27946 Dr.28581 OPHN1 Transcribed locus Oligophrenin Dr.14931 Dr.81311 ZGC:153327 DRL Dr.76198 Dr.75142 Dr.86287 IM:7150932 SI:CH211-197G15.1 ZGC:63651 protein Zgc:153327 Hypothetical protein LOC791603 Im:7150932 Si:ch211-197g15.1 Zgc:63651 255 [...]... Lgr5 not only marks stem cells in the small intestine, but also marks stem cells in the hair follicle [48] It is also suggested to mark stem cells in the stomach and mammary gland [34, 45] Like Lgr5, Ascl2 has also been identified to mark intestinal stem cells in a more specific manner [36] Transcriptome of intestinal stem cells have been probed by microarray [36] and culture of the stem cells has been reported... understanding of the nature of the intestinal biology 1.5.2 Intestinal stem cell number The number of intestinal stem cells is under tight regulation under normal physiology Excessive stem cells are believed to cause crypt division or fission, thereby maintaining the desired number of stem cells within each crypt Stem cells are sensitive to irradiation and irradiation-induced DNA damage will cause the stem. .. same source of stem cells [18] Following that, increasing attention has been drawn to this field of research due to the potential medical importance [19] Evidence 8 in support of the existence of intestinal stem cells and their multipotency has been ever growing 1.5.1 Location of intestinal stem cells Historically, there have been two schools of thought regarding the location of intestinal stem cells One... mouse intestine [34, 35, 36] Proof of pluripotency of the intestinal stem cells based on lineage tracing in genetically modified mice has been equally successful in support of either school of thoughts [28, 37, 34, 36] Current results, therefore, appear to support the presence of two populations of intestinal stem cells, which differ in their location, cell number and molecular profiles In view of their... factor, and the antibacterial cryptins [11] 1.3 Turnover of the intestinal epithelium The intestinal epithelium represents one of the most rapidly renewing tissues of the human body [6] The epithelial cells of the intestine undergo apoptosis at the tips of villi, and the sloughed cells are replaced by neighboring cells migrating upward In the crypts, new cells are generated by stem/ progenitor cells to... damage will cause the stem cells to undergo apoptosis in an altruistic manner to prevent passing the DNA damage to their daughter cells Loss of stem cells will be compensated by expansion of the remaining stem cells or their immediate daughter cells that still possess stemness property or with colonogenic potential Due to unavailability of molecular makers, the number of stem cells has not been verified... position of the crypt and these cells are able to generate the four major epithelial cell types (columnar cells, goblet cells, enteroendocrine cells and Paneth cells) in mouse small intestine [28] 9 The second school believes that the stem cells are sandwiched between the post-mitotic Paneth cells near the bottom of the crypts based on the identification of crypt base columnar (CBC) cells, which are small,... absence of villous structures in frog intestine till the metamorphic stage [67, 68] Following the metamorphosis, adult frog intestine features presence of both villi and crypts (also called crests and troughs, respectively) The intestinal epithelium contains columnar cells, goblet cells and enteroendocrine cells [69, 70] The columnar cells are most abundant and the goblet cells take up about 16 10% of the... and molecular aspects Features of small intestine and large intestine will be explored and molecular similarity between regions of zebrafish intestine and their mammalian counterparts will be investigated Research goals of the thesis work are briefly outlined in the next section 1.8 Research goals of the current work The current thesis work aims to have a better understanding of the regionalization and. .. Around 14 dpf, degenerating cells appear at the top of the epithelial elevations These cells become rounded and are extruded into the lumen Adult mouse features presence of crypts and villi in the small intestine, but only crypts in the large intestine [34] Each crypt contains a monoclonal popu- 13 lation of cells [50, 51] derived from multipotent stem cells [52] After a phase of rapid amplification, the . INTESTINE OF ZEBRAFISH: REGIONALIZATION, CHARACTERIZATION AND STEM CELLS WANG ZHENGYUAN (M.Sci., NUS) (B.Eng., NPU) A THESIS SUBMITTED FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN COMPUTATION AND. characteristics of the crypt-villus system . . . . 158 5.3.2 Determination of the number of epithelial stem cells in a 2D section of the inter-villi pocket of zebrafish (Danio rerio) intestine . number of stem cells is about 2 to 4 per section of an inter-villi pocket in the small intestine of zebrafish. Interestingly, this number seems to remain similar in the small intestines of other