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The above model is well illustrated by consideration of the astrocytoma Among the rst detectable changes are mutations that inactivate the tumor suppressor gene, p53 on chromosome 17p; over 50 percent of astrocytomas have deletions encompassing this gene Other early changes include overexpression of growth factors or their receptors as noted below After the tumor develops, progression to a more malignant grade of astrocytoma or to a glioblastoma may be triggered by defects in the p16-retinoblastoma gene signaling pathway, loss of chromosome 10 (seen in about 90 percent of high-grade gliomas), or overexpression of the epidermal growth factor gene In fact, it is striking that analysis of the patterns of these defects correlates accurately with the staging and aggressive characteristics of these tumors Knowledge of the molecular signatures of certain other tumors has immense clinical value For example, as discussed further on, oligodendrogliomas that have combined deletions in chromosomes 1p and 19q respond well to chemotherapy, and this property may increase survival by more than 10 years This type of information may spare the nonresponsive patient from ineffective, sometimes toxic therapy (see Reifenberger and Louis; Louis et al) Much of the modern genetic classi cation of brain tumors is derived from the technical tour de force of gene microarrays The patterns of these multiple gene analyses are able to distinguish some types of medulloblastomas from the similar-appearing primitive neuroectodermal tumors; the medulloblastomas express classes of genes that are characteristic of cerebellar granule cells, suggesting they arise from these cells Also, these gene expression signatures confer useful prognostic information in a more general way than noted above for oligodendroglioma For example, medulloblastomas that express genes indicative of cerebellar differentiation are associated with longer survival than those expressing genes related to cell division (Pomeroy et al) Furthermore, as alluded to earlier, in about 50 percent of gliomas there is an overexpression or a mutant form of epidermal growth factor (EGF) and of transforming growth factor alpha (TGF- ), suggesting a role for these in the progression of certain tumor types Other trophic factors are overexpressed in yet other brain neoplasms and perhaps contribute to their morphology and growth pattern; for example, VEGF is found in extremely high concentrations in meningiomas, which are highly vascular by nature These ndings, taken together, suggest an autocrine stimulation of growth by these factors and possibly an interaction with some of the aforementioned gene defects However, having emphasized molecular and chromosomal changes, it is not yet clear if any of them is truly causative (the currently favored hypothesis) or if they simply re ect an aberrant genetic process that accompanies the dedifferentiation of tumor growth and progression On the basis of this new molecular information, our views of the pathogenesis of neoplasia are being cast along new lines Some of the speci cs of these new data are presented in the following discussions of particular tumor types A more extensive review can be found in the article by Osborne and colleagues, including the heterogeneity of ndings that suggests polygenic changes in most gliomas.

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