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DEVELOPMENTAL CONTROL OF HUMAN GLOBIN GENES George Stamatoyannopoulos, M.D., Dr i. University of Washington Seattle, Washington USA Investigations aimed to understand the cellular and molecular mechanisms that control hemoglobin switching during development span almost one-half of a century. Special emphasis has been given to the understanding of the control of the synthesis of fetal hemoglobin in adult life and several studies have focused on the development of therapeutic modalities for induction of fetal hemoglobin synthesis in adult individuals. Research done in the 1970s led to insights on the cellular control of fetal hemoglobin synthesis in adult individuals and provided evidence that fetal hemoglobin can be induced in the adult by several manipulations that result in stressed erythropoiesis. These developments led to the introduction of cytotoxic drugs for induction of fetal hemoglobin and the development of hydroxyurea as a therapeutic modality in sickle cell syndromes. Systematic investigations of the molecular mechanisms of hemoglobin switching started in the 1980s, when new molecular technologies for studying the control of gene expression were introduced. A seminal contribution to the understanding of the molecular control of globin gene expression was the discovery in the late 80s of the globin locus control region. This discovery stimulated a large number of investigations and greatly facilitated experimentation using transgenic mice. Another methodological development has been the use of YACs in the analysis of hemoglobin switching. Several studies in transgenic mice have shown that two mechanisms, autonomous silencing and gene competition, control hemoglobin switching. During the last several years, my group has focused on the investigation of the mechanisms by which the fetal globin genes are turned on and off during the course of development. Experimental evidence indicates that turning on of the g globin genes involves interactions between the locus control region and the globin gene promoter. We have identified the sequences of the LCR and sequences of the globin gene promoter involved in these interactions, but the transcriptional factors that mediate these interactions are still unknown. There is strong evidence that the globin genes are turned off by autonomous silencing and that all the sequences participating in gene silencing are located in the globin gene promoter. gene silencing most likely is the result of disruption of the interaction between the globin gene promoter and the LCR, but the trans-acting elements that mediate this phenomenon are unknown. We postulate that gene silencing takes place during the course of erythroid differentiation in the adult erythropoiesis and that escape from gene silencing is the main mechanism whereby fetal hemoglobin continues to be expressed in a subpopulation of erythroid cells of the adult. Inhibition of silencing is the most likely mechanism of action of fetal hemoglobin inducers currently in use and represents the most useful approach for the development of new inducers of fetal hemoglobin in patients with chain hemoglobinopathies and mobic.
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We have audited the Financial statements which comprise the consolidated statement of profit and loss, consolidated statement of total recognised gains and losses, consolidated statement of cash flow, consolidated and company balance sheets and the related notes, which have been prepared under the historical cost convention and the accounting policies set out in the statement of accounting policies. We have also audited the disclosures required by Part 3 of Schedule 7A to the Companies Act 1985 contained in the Directors' Remuneration Report `the auditable part' ; . Respective responsibilities of Directors and Auditors The Directors' responsibilities for preparing the annual report and the Financial statements in accordance with applicable United Kingdom law and accounting standards are set out in the statement of Directors' responsibilities. The Directors are also responsible for preparing the Directors' Remuneration Report. Our responsibility is to audit the Financial statements and the auditable part of the Directors' Remuneration Report in accordance with relevant legal and regulatory requirements and United Kingdom Auditing Standards issued by the Auditing Practices Board. This report, including the opinion, has been prepared for and only for the company's members as a body in accordance with Section 235 of the Companies Act 1985 and for no other purpose. We do not, in giving this opinion, accept or assume responsibility for any other purpose or to any other person to whom this report is shown or into whose hands it may come save where expressly agreed by our prior consent in writing. We report to you our opinion as to whether the Financial statements give a true and fair view and whether the Financial statements and the auditable part of the Directors' Remuneration Report have been properly prepared in accordance with the Companies Act 1985. We also report to you if, in our opinion, the Directors' Report is not consistent with the Financial statements, if the company has not kept proper accounting records, if we have not received all the information and explanations we require for our audit, or if information specified by law regarding Directors' remuneration and transactions is not disclosed. We read the other information contained in the Annual Report and consider the implications for our report if we become aware of any apparent misstatements or material inconsistencies with the Financial statements. The other information comprises only the Financial summary, description of business, unauditable part of the Directors Remuneration Report, the joint statement by the Chairman and the Chief Executive Officer, the operating and financial review and prospects and the corporate governance statement. We review whether the corporate governance statement reflects the company's compliance with the nine provisions of the 2003 FRC Combined Code specified for our review by the Listing Rules of the Financial Services Authority, and we report if it does not. We are not required to consider whether the Board's statements on internal control cover all risks and controls, or form an opinion on the effectiveness of the company's or Group's corporate governance procedures or its risk and control procedures and protonix.
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