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A Systems-Based Approach for the Characterization of Toxicity Pathways Associated with the HPG Axis A Genomics Timeline © 2003 Nature Publishing Group DESIGN BY DARRYL LEJA PEAS COURTESY J. BLAMIRE, CITY UNIV. NEW YORK; WATSON & CRICK COURTESY A. BARRINGTON BROWN/SPL; SCIENCE COVERS COURTESY AAAS (Toxico) Genomics: An Emerging Science* 200 150 2000 100 1000 50 0 0 95 96 97 98 99 00 01 02 03 04 05 Year Current Contents, 10/05 Toxi cogenom i cs Publ i cat i ons ( ) G enomi cs Publ i cat i ons( ) 3000 Biological Responses and Genomics: An Overview WHAT RESPONSES ARE POSSIBLE GENOMICS INITIATION OF RESPONSE TRANSCRIPTOMICS WHAT DRIVES THE RESPONSE Adapted from Viant (2005) PHENOTYPE PROTEOMICS THE METABOLIC RESPONSE METABOLOMICS Transcriptomics: Example of a Microarray Experiment Study Goal: Examine fadrozole MOA using a 2,000 gene fathead minnow oligonucleotide array • Multiple tissues examined • Multiple differentially-regulated genes Spot color Regulation Up-regulated genes Down-regulated genes No change Data from EPA/ EcoArray© CRADA Proteomics: Example of a Gel Separation/MALDI-TOF MS Experiment Intens. [a.u.] Peptide Mass Fingerprinting x10 4 1091.620 1.5 1799.879 1347.669 1.0 Representative protein expression profile in testes of control zebrafish 2143.156 1615.722 890.612 1214.658 0.5 1504.667 1978.039 2460.281 2801.340 0.0 1000 1500 2000 Data from EPA-Cincinnati 2500 3000 m /z Metabolomics: Example of an NMR Experiment Fathead Minnow (male) Sprague-Dawley Rat (male) Data from EPA-Athens Toxicogenomics: Benefits • Unprecedented ability to generate a global “picture” of the health status of organisms • Offers unique potential to define MOA/toxicity pathways from two perspectives – Identification of key molecular targets/control nodes – Linkage of responses across biological levels of organization • Both perspectives critical to successful ecological risk assessments Toxicogenomics: Challenges • Amount of data to consider is daunting, well exceeding historical ecotoxicology bioinformatic capabilities – Example: ~30,000 genes/100,000 proteins/20,000(?) metabolites (humans) • Identity of many (sometimes majority) of altered genes/proteins/metabolites uncertain in key species for ecotoxicology research • Even well-defined treatments can cause many changes, complicating interpretation of biological significance – Example: zebrafish exposed to fadrozole - up to 1000 unique up - or down - regulated genes in brain and gonad (21K microarray) Conceptual Systems Models • Can help focus data analysis on relevant genes/proteins/metabolites • Provide an a priori framework for formal hypothesis testing of observed changes • Allow “discovery” of unanticipated system components/control nodes key to biological responses • Iterative testing/model modification establishes basis for predictive computational model (s) 1 2 3 4 5 6 7 Brain 8 9 10 11 12 13 14 15 16 18 19 20 Figure Key state transition a Catalysis (including Liver activation) transcriptional activation b translational activation transcription inhibition c dissociation association d Genes mRNA e protein activated f protein g receptor Simple h molecule Phenotype i j Pituitary k Blood l m n o p Ovary q r Graphical Systems Model for Small Fish HPG Axis Characteristics of Conceptual HPG System • Constructed using Cell Designer® 3.1 coded in SBML • Captures 7 functional modules in 6 tissue compartments with multiple subcompartments (cell types) for brain, pituitary, ovary and testis • Assembled from more than 60 primary literature and review papers on vertebrate HPG axis • Depicts interactions of over 105 proteins, 40 simple molecules, regulation of 25 genes, and over 300 reactions relative to our current understanding of the system Linkage of Exposure and Effects Using Genomics, Proteomics, and Metabolomics in Small Fish Models • USEPA – Cincinnati, OH – D. Bencic, I. Knoebl, D. Lattier, J. Lazorchak, G. Toth, R. Wang • USEPA – Duluth, MN, and Grosse Isle, MI – G. Ankley, E. Durhan, K. Jensen, M. Kahl, L. Makynen, D. Martinovic, D. Miller, D. Villeneuve • USEPA – Athens, GA – T. Collette, D. Ekman, T. Whiteside • USEPA-RTP, NC – M. Breen, R. Conolly • USEPA STAR Program – N. Denslow (Univ. of Florida), E. Orlando, (Florida Atlantic University), K. Watanabe (Oregon Health Sciences Univ.), M. Sepulveda (Purdue Univ.) • USACOE - Vicksburg, MS – E. Perkins • DOE Partners – Joint Genome Institute, (Walnut Creek, CA) – Sandia, (Albuquerque, NM) – PNNL (Richland, WA) Compartment GABA Dopamine Brain ? ? ? PACAP Pituitary GnRH Neuronal System GnRH NPY D2 R GABAB R Y2 R GnRH R 2 Muscimol (+) 3 Apomorphine (+) 4 Haloperidol (-) Y1 R Gonadotroph Activin R Fipronil (-) Y2 R GABAA R PAC1 R 1 D1 R Follistatin Activin Chemical “Probes” D2 R GPa FSHb Blood Circulating LDL, HDL LHb Circulating LH, FSH LDL R LH R 5 Trilostane (-) 6 Ketoconazole (-) FSH R HDL R Cholesterol Outer mitochondrial membrane StAR Inner mitochondrial membrane Gonad Activin (Generalized, gonadal, steroidogenic cell) Inhibin P450scc pregnenolone 3bHSD 17α-hydroxyprogesterone Fadrozole (-) 8 Prochloraz (-,-) progesterone P450c17 20βHSD 7 androstenedione 17βHSD 17α,20β-P (MIS) testosterone P450arom 9 P45011β. Vinclozolin (-) 11βHSD 11-ketotestosterone Blood Androgen / Estrogen Responsive Tissues (e.g. liver, fatpad, gonads) Circulating Sex Steroids / Steroid Hormone Binding Globlulin ER AR 10 Flutamide (-) 11 β-trebolone (+) 12 Ethynyl estradiol (+) estradiol Effects of Aromatase Inhibition on Reproduction in the Fathead Minnow 150 N 10 N a Aromatase Activity (fmol/mg-1 hr-1) 2 10 6 Male Female Control 50 * * * 4 Fadrozole 75 c 2 c CN 0 -20 -18 -16 -14 -12 -10 -8 -6 -4 -2 0 2 4 6 8 10 12 14 16 18 20 0 Exposure (d) 0 6 E2 (ng/ml) 50 Fadrozole (µg / L) 8 4 * 2 * 0 Vtg (mg/ml) Cumulative Number of Eggs (Thousands) Fadrozole (ug/L) 8 b 20 10 * * 0 Control 2 10 Fadrozole (µg/l) * 50 Fadrozole Genomic Analyses • Fathead minnows exposed to varying concentrations of fadrozole for 24h or 7d • Liver, brain and gonad collected and gene expression assessed by microarray or PCR • Depending upon statistical criteria, 10s to 100s of genes changed in each tissue of females exposed for 7 d • Interpretation questions – Meaningful to biological function? – Significant to HPG axis? – Direct versus compensatory response(s)? Gene Expression in Fadrozole-Treated Fathead Minnows: Hypothesized vs. Observed Changes1 Hypothesized Observed (Fold)2 - - Vtg precursor - (18-70) -Vtg 3 precursor - (7-98) Zrp NC3 ERa (2.0-2.8) EST NP HSST NP CYP3A (1.6) Gene Vtg 1 Livers from females exposed for 7 d 2 NC, no change; NP, no probe for gene on microarray Changes in HPG-Relevant Genes in Fathead Minnows Exposed to Fadrozole1 Gene Observed (Fold) Aromatase B (9.6) Cyclin B (5.8-7.2) Cytochrome B (13-170) GAD 65 (1.2-2.6) GAD 67 (1.8-2.6) HMG-CoA Reductase (2.8-5.4) IGF Binding Protein (220) LDLR Associated Protein (1-16) Pit-1 (9.4) PRMC (2.1-5.3) 1 Ovaries from females exposed for 7-d Fadrozole Paracrine Inner mitochondrial membrane transport Granulosa cell Theca cell Linkages Across Biological Levels of Organization: Toxicity "Pathways" for HPG-Active Chemicals Molecular • Gene/Protein Expression • Metabolite Profiles Cellular Alterations in production of signaling molecules Organ • Functional changes • Structural changes (pathology) Individual Population Altered reproduction or development Decreased numbers of animals Increasing Ecological Relevance Increasing Diagnostic (Screening) Utility Linkage of Molecular Responses to Population Effects • Molecular responses (biomarkers) need to reflect toxicity pathways of concern • Molecular responses also require biologically plausible (ideally quantitative) linkages to adverse outcomes in individuals • Individual outcomes need to be easily translated into ecologically-realistic population context Vitellogenin as a Biomarker: Linkage to Population-Level Effects • Decreases in Vtg in females are a consistent (and mechanistically reasonable) response to decreases in steroidogenesis caused by chemicals • Vtg status also has been hypothesized as a direct indicator of female’s ability to produce eggs • If the relationship “holds”, population-level responses based on Vtg should be possible Lab Test Data Life Table - Leslie Matrix Carrying Capacity/Habitat Quality Density-Dependent Model for Population Prediction Model Application and Results Fecundity in Fecundity Decrease Measurement of vtg concentrations and fecundity for female fathead minnows 17β-trenbolone Projection of density dependent logistic population trajectories for the fathead minnow population based upon change in vtg 17α-trenbolone prochloraz prochloraz fenarimol fenarimol fadrozole fadrozole DecreaseVtg in VTG Chemical Life table with age specific vital rates of survival and fecundity for the fathead minnow population Carrying capacity for the fathead minnow population Fathead Minnow Fecundity vs Vtg Exposure Concentrations 1 0.005µg/l, 0.05µg/l, 0.5µg/l, 5µg/l, and 50µg/l 0.003µg/l, 0.01µg/l, 0.03µg/l, and 0.1µg/l 0.03mg/l, 0.1mg/l, and 0.3mg/l 0.1mg/l and 1mg/l 2µg/l, 10µg/l, and 50µg/l Fecundity = -0.042 + 0.95 * Vtg (R2 = 0.88) 0.9 0.8 0.7 Relative Fecundity 17β-trenbolone 17α-trenbolone Prochloraz Fenarimol Fadrozole 0.6 0.5 0.4 0.3 0.2 0.1 0 0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 Relative Vitellogenin 0.8 0.9 1 Model Application and Results Measurement of vtg concentrations and fecundity for female fathead minnows Fecundity 17β-trenbolone Projection of density dependent logistic population trajectories for the fathead minnow population based upon change in vtg 17α-trenbolone prochloraz fenarimol fadrozole Vtg Life table with age specific vital rates of survival and fecundity for the fathead minnow population Carrying capacity for the fathead minnow population Population projection for populations at carrying exposed to stressors that depress vitellogenin production Average Population Size Size Average Population (Proportion of Carrying Capacity) (Proportion of Carrying Capacity) Forecast Population Trajectories 1 1 A A 0% 0.8 0.8 0.6 0.6 0.4 0.4 B B 25% 0.2 0.2 E D D 0 >95%E 75% 0 C C 50% 0 0 5 5 10 10 Time (Years) Time (Years) 15 15 20 20 It’s tough to make predictions, especially about the future. Yogi Berra New York Philosopher