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Copyright =C2=A9 2004=20 Cell Press.

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Getting at the Root and Stem of Brain Tumors

Trudy G. Oliver and Robert J. Wechsler-Reya,
Department of Pharmacology and Cancer = Biology,=20 Duke University Medical Center, Durham, NC 27710, USA

Available = online=20 23 June 2004.

Abstract

Brain tumors are among the most aggressive and intractable types of = cancer.=20 Recent studies indicate that brain tumor cells resemble neural stem = cells in=20 terms of phenotype, signaling, and behavior in vitro. In light of these=20 similarities, it has been suggested that brain tumors arise from stem = cells,=20 that they co-opt stem cell strategies for self-renewal, and even that = they=20 contain =E2=80=9Ccancer stem cells=E2=80=9D that are critical for tumor = maintenance. We will=20 examine these possibilities and discuss their implications for the = understanding=20 and treatment of brain tumors.


Article Outline

=E2=80=A2=20 Main Text
=E2=80=A2=20 Genes and Signals Shared by Brain Tumors and Stem Cells
=E2=80=A2=20 Self-Renewal and Multipotency of Brain Tumors and Stem Cells
=E2=80=A2=20 Do Brain Tumors Arise from Neural Stem Cells?
=E2=80=A2=20 Do Brain Tumors Contain =E2=80=9CCancer Stem = Cells=E2=80=9D?
=E2=80=A2=20 References

Main Text

Due to their rapid growth and tendency to spread throughout the brain = and=20 spinal cord, brain tumors are extremely difficult to treat and often = fatal. Of=20 the 18,000 Americans diagnosed with primary malignant brain tumors each = year,=20 only one-third will survive beyond 5 years (American=20 Cancer Society, 2004). Glioblastoma multiforme, the most common = brain tumor=20 in adults, has a mean survival time of 10=E2=80=9312 months. = Medulloblastoma, the most=20 common malignant brain tumor in children, has a more favorable = prognosis:=20 combinations of radiation, chemotherapy, and surgery can cure more than = half of=20 patients. However, children who survive medulloblastoma treatment often = suffer=20 severe cognitive and physical deficits and increased susceptibility to = other=20 cancers. To develop more effective therapies for these tragic diseases, = we need=20 a deeper understanding of their molecular and cellular basis. Recent = studies=20 suggest that this understanding may come from an appreciation of the=20 relationship between brain tumors and neural stem cells (NSCs).

Genes and Signals Shared by Brain Tumors and Stem = Cells

The fact that brain tumors frequently include a mixture of neuronal = and glial=20 cell types has often been cited as evidence that these tumors may = contain=20 multipotent progenitors. Consistent with this notion, many brain tumors = contain=20 cells that express markers of neural stem cells (Hemmati=20 et al. 2003; Ignatova=20 et al. 2002; Leung=20 et al. 2004 and Singh=20 et al. 2003). The most common example of a shared antigen is the=20 intermediate filament protein nestin, which is expressed in NSCs in the=20 developing and adult CNS and has also been found in brain tumors and = brain=20 tumor-derived cell lines. Recent studies have reinforced this = correlation by=20 describing expression of other NSC markers=E2=80=94including CD133, = musashi-1,=20 bmi-1, and sox-2=E2=80=94in primary brain tumor cells. = Although the=20 phenotypic similarities between brain tumors and NSCs could be = coincidental, the=20 growing list of commonly expressed genes suggests a more intimate = relationship=20 between these cells.

In fact, the similarity between brain tumors and neural stem cells = extends to=20 the level of cell signaling as well. For example, the Sonic hedgehog = (Shh)=20 pathway plays a critical role in regulating growth of neural progenitors = and has=20 been implicated in the etiology of brain tumors. In the cerebellum, Shh = is=20 required for proliferation of granule cell precursors, and mutations in = this=20 pathway predispose to cerebellar tumors (medulloblastomas) in both mice = and=20 humans (reviewed in Wechsler-Reya,=20 2003). Recent studies suggest that Shh signaling may also regulate=20 self-renewal of progenitors in other parts of the brain, including the=20 subventricular zone, hippocampus, and olfactory bulb (Lai=20 et al. 2003 and Machold=20 et al. 2003). These findings raise the possibility that Shh pathway=20 activation may be involved in tumors besides medulloblastoma. Although a = small=20 percentage of glial tumors (gliomas) exhibit amplification of the = Shh-responsive=20 transcription factor Gli1, no other Shh pathway mutations have = been found=20 in these tumors. However, at least one report suggests that hedgehog = pathway=20 genes are expressed in gliomas and that growth of glioma cell lines is = reduced=20 by pharmacological inhibitors of the pathway (Dahmane=20 et al., 2001). In light of recent evidence that ligand-dependent = (and=20 mutation-independent) hedgehog pathway activation contributes to = tumorigenesis=20 in other tissues (Berman=20 et al., 2003), the role of Shh signaling in other brain tumors = merits=20 further study.

The Wnt pathway, which is critical for self-renewal of hematopoietic = and=20 epithelial stem cells (Reya=20 et al., 2001), may also be involved in the growth of NSCs and brain = tumors.=20 Recent reports show that =CE=B2-catenin, a central mediator of Wnt = signaling, is=20 important for NSC and progenitor cell proliferation throughout the CNS.=20 Conditional deletion of =CE=B2-catenin causes depletion of = neuronal precursors=20 in the ventricular zone, whereas overexpression of activated = =CE=B2-catenin results=20 in massive expansion of the precursor pool and enlargement of the = hindbrain,=20 midbrain, and forebrain (Chenn=20 and Walsh 2002 and Zechner=20 et al. 2003). Activation of the Wnt pathway is also associated with = brain=20 tumors. In particular, a subset of medulloblastomas has been reported to = harbor=20 mutations in =CE=B2-catenin, Axin, or APC (Wechsler-Reya,=20 2003). Since these tumors have a distinct phenotype and molecular = profile=20 from medulloblastomas associated with hedgehog pathway mutations, they = may arise=20 from a distinct set of progenitors. To date, there have been no reports = of Wnt=20 pathway mutations in brain tumors outside the cerebellum. However, given = the=20 importance of Wnt signaling in self-renewal of progenitors throughout = the brain,=20 it is worth considering whether dysregulation of this pathway occurs in = other=20 types of brain tumors as well.

Other genes that may control self-renewal of both brain tumors and = neural=20 stem cells include the polycomb transcription factor Bmi-1 and the = phosphatase=20 PTEN. Bmi-1 regulates self-renewal of progenitors in the central and = peripheral=20 nervous systems (Molofsky=20 et al., 2003), in part by repressing genes (such as p16Ink4a and = p19Arf)=20 that promote cellular senescence and cell death. In addition, = bmi-1 has=20 recently been shown to be a target of the Shh signaling pathway and to = be=20 required for self-renewal of neural progenitors in the cerebellum (Leung=20 et al., 2004). Thus, elevated expression of bmi-1 in = medulloblastoma=20 and other brain tumors (Hemmati=20 et al. 2003 and Leung=20 et al. 2004) may reflect an increased capacity for self-renewal. = PTEN=20 is one of the most frequently mutated genes in primary glioblastoma (Li=20 et al., 2003). Recent studies of mice in which PTEN has been=20 conditionally deleted in nestin-expressing precursors suggest that it is = a=20 critical negative regulator of NSC proliferation and survival (Groszer=20 et al., 2001). In addition, studies of = PTEN+/=E2=88=92 mice=20 suggest that this gene may also be important for NSC migration (Li=20 et al., 2003). Thus, dysregulation of PTEN may contribute to the = aggressive=20 growth and metastatic behavior associated with gliomas.

Self-Renewal and Multipotency of Brain Tumors and Stem = Cells

The similarities in phenotype and signaling between brain tumor cells = and=20 stem cells may reflect similarities in their cellular behavior. Support = for this=20 notion comes from recent studies by Ignatova=20 et al. (2002), Hemmati=20 et al. (2003), and Singh=20 et al. (2003). These investigators isolated cells from primary human = brain=20 tumors (including astrocytomas, glioblastomas, and medulloblastomas) and = showed=20 that, like NSCs, they form clonal neurospheres that can be repeatedly = passaged=20 in vitro. Upon withdrawal of growth factors, tumor-derived neurospheres = can=20 differentiate into both neurons and glia, consistent with the = heterogeneous cell=20 types found in the tumors from which they were derived. Although brain = tumors=20 have long been known to contain a mixture of neurons and glia, it has = been=20 unclear whether all of these cells are derived from the tumor or whether = some=20 represent normal cells that have become trapped within the tumor matrix. = The=20 observation that brain tumor-derived neurospheres can generate multiple = cell=20 types suggests that the heterogeneity within brain tumors may be an = inherent=20 characteristic of the tumor. Together, these studies suggest that brain = tumor=20 cells share with neural stem cells the capacity for self-renewal and=20 multilineage differentiation.

In interpreting these studies, it is important to consider the = possibility=20 that prolonged culture of progenitors in high concentrations of growth = factors=20 may cause cells to adopt fates that they would not normally adopt in = situ, or=20 select for a subset of cells that does not reflect the behavior of the = original=20 population (Gabay=20 et al., 2003). Thus, culture of brain tumor cells under=20 neurosphere-generating conditions may dramatically alter their capacity = for=20 self-renewal and differentiation. Studying tumor cells that have not = been=20 subjected to extensive culture in growth factors may help shed light on = their=20 intrinsic properties. Notwithstanding these caveats, the demonstration = of=20 functional similarities between brain tumor cells and NSCs represents an = important step beyond phenotypic characterization and brings us closer = to=20 understanding how these cells may be related to one another.

Do Brain Tumors Arise from Neural Stem Cells?

One explanation for the resemblance between brain tumors and stem = cells is=20 that brain tumors result from transformation of NSCs (see Figure=20 1). But does the fact that brain tumor cells look and act like NSCs = mean=20 that they actually arise from them? Or can they result from acquisition = of stem=20 cell characteristics by more differentiated progenitors? There is = evidence for=20 both possibilities.=20

Display=20 Full Size version of this image = (15K)

Figure 1.=20

The Relationship between Brain Tumors and Neural Stem Cells

Brain tumors (red) share many characteristics with NSCs (yellow), = including=20 expression of intracellular and cell surface markers, activation of = signaling=20 pathways, and an ability to form self-renewing, multipotent = neurospheres in=20 vitro. One explanation for these similarities is that brain tumors = result from=20 transformation of NSCs, and there is evidence from transgenic mice to = support=20 this possibility. However, brain tumors may also arise from = transformation of=20 more restricted progenitors; in this case, the similarities in gene = expression=20 might represent acquisition of stem cell characteristics as a = consequence of=20 transformation. In either case, cells with stem cell-like properties = are=20 likely to be critical for growth of brain tumors, and targeting these = cells=20 may represent an important avenue for therapy.=20

Support for the notion that brain tumors can arise from = transformation of=20 neural stem cells comes from mouse models in which expression of = oncogenes or=20 tumor suppressors is perturbed in an NSC-specific manner. For example, = Holland=20 and colleagues have generated mice that express the avian retrovirus = receptor=20 (TVA) in nestin+ cells, and then used avian retroviruses to deliver = oncogenes=20 into neural stem cells (reviewed in Holland,=20 2001). Using this system, they have shown that transduction of=20 constitutively active epidermal growth factor receptor, oncogenic Ras = and Akt,=20 or platelet-derived growth factor into forebrain nestin+ progenitors = results in=20 tumors that resemble human gliomas. Similarly, intracerebellar injection = of=20 retroviruses carrying shh and c-myc can cause = medulloblastoma in=20 nestin-TVA mice (Rao=20 et al., 2003). To the extent that transgene expression in these mice = is=20 restricted to neural stem cells (nestin expression has also been = observed in=20 more committed progenitors), these studies suggest that NSCs can serve = as=20 targets of transformation in brain tumors.

But there is also evidence that brain tumors can result from = transformation=20 of more differentiated cells. Shh pathway-associated medulloblastomas in = both=20 mice and humans have a phenotype and gene expression profile that = strongly=20 resembles that of committed granule cell precursors (Wechsler-Reya,=20 2003). Moreover, in studies using the TVA system, gliomas can be = generated=20 by retroviral transduction of genes into GFAP+ glial progenitors as well = as=20 nestin+ NSCs (Holland,=20 2001). Expression of oncogenic Ras or SV40 T antigen under the = control of a=20 GFAP promoter also results in gliomas in a large percentage of = animals (Ding=20 et al. 2001 and Xiao=20 et al. 2002). Finally, transgenic mice expressing v-erbB driven by = an=20 S100-=CE=B2 promoter (which is active in glial precursors) = develop=20 oligodendrogliomas (Weiss=20 et al., 2003). These studies strongly suggest that restricted = progenitors=20 may be targets for transformation by oncogenes. However, since GFAP and = S100=CE=B2=20 may also be expressed in neural stem cells (Goldman=20 2003 and Weiss=20 et al. 2003), it will clearly be necessary to identify more = definitive=20 markers of stem cells and progenitors to determine the cell of origin of = brain=20 tumors.

Do Brain Tumors Contain =E2=80=9CCancer Stem = Cells=E2=80=9D?

The presence of NSC-like cells in brain tumors has also been used to = suggest=20 that these tumors contain =E2=80=9Ccancer stem cells=E2=80=9D that are = critical for their=20 propagation (Hemmati=20 et al. 2003 and Singh=20 et al. 2003). While this is an intriguing possibility, it is = critical to=20 stress the distinction between tissue stem cells (such as NSCs) and = cancer stem=20 cells (Reya=20 et al., 2001). Tissue stem cells exhibit extensive self-renewal and = have the=20 capacity to differentiate into multiple cell types in the normal tissue = from=20 which they are derived (e.g., neurons, astrocytes, and oligodendrocytes = in the=20 CNS). Cancer stem cells may also exhibit extensive self-renewal and=20 multipotency. But most importantly, cancer stem cells are defined by = their=20 capacity to maintain the long-term growth of a tumor in vivo. This = capacity for=20 tumor propagation is most compellingly demonstrated by transplanting = putative=20 cancer stem cells into a new host and showing that they can recapitulate = the=20 growth of the tumor. Such experiments have been carried out with human = leukemia=20 and breast cancer cells and have shown that a very small fraction of the = original tumor is uniquely capable of recapitulating the tumor following = transplant (Al-Hajj=20 et al. 2003 and Bonnet=20 and Dick 1997).

In the case of brain tumors, Singh et al. show that a subset of brain = tumor=20 cells (those expressing CD133) have the exclusive ability to form = multipotent=20 neurospheres in vitro. While isolation of this population is an = important step,=20 the ability of these cells to generate and maintain tumors in vivo = remains to be=20 tested. If CD133+ cells (or any other population within these tumors) = represent=20 true cancer stem cells, it would have significant implications for the=20 understanding and treatment of brain tumors. First, it would suggest = that=20 despite the heterogeneity inherent in many brain tumors (reflected, for = example,=20 by the term glioblastoma multiforme), a single cell type may be=20 responsible for tumor growth and maintenance. Identification of this = cell type=20 might provide important markers that can be used for tumor detection and = diagnosis. Finally, targeting of this cell may be necessary (and perhaps = even=20 sufficient) to disrupt the growth of the tumor.

The studies described above suggest that brain tumors and neural stem = cells=20 resemble one another at a cellular and molecular level. Whether brain = tumors=20 actually arise from NSCs in most cases remains unclear, and a definitive = answer=20 may have to await the development of better markers for neural stem = cells and=20 progenitors. Nevertheless the fact that these cells express similar = genes and=20 show activation of similar signaling pathways suggests that they use = similar=20 strategies to achieve self-renewal and to generate heterogeneous = populations of=20 cells within a tissue. Understanding stem cell signaling pathways and = developing=20 ways to manipulate them will be critical in our effort to develop new = treatments=20 for tumors of the nervous system.


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Corresponding author. Correspondence: Robert J.=20 Wechsler-Reya, (919) 613-8754 (phone), (919) 668-3556 (fax) =



Volume=20 42, Issue 6, 24 June 2004, Pages 885-888
 
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1px solid; WIDTH: 270px; = BORDER-BOTTOM: #cccccc 1px solid; HEIGHT: 100px; BACKGROUND-COLOR: = #ffffff } .mlktLink { PADDING-RIGHT: 0px; PADDING-LEFT: 0px; PADDING-BOTTOM: 5px; = PADDING-TOP: 5px; BACKGROUND-COLOR: #ffffff } .mlktEmpty { PADDING-RIGHT: 5px; PADDING-LEFT: 5px; PADDING-BOTTOM: 5px; = PADDING-TOP: 5px; BACKGROUND-COLOR: #ffffff } ------=_NextPart_000_0000_01C8569C.7BCDC930 Content-Type: text/css; charset="iso-8859-1" Content-Transfer-Encoding: quoted-printable Content-Location: http://www.sciencedirect.com/science/page/static/stylesheets/css_gen_v01.css BODY { MARGIN: 0.4em 0em 0em } DIV { FONT-SIZE: 1em; FONT-FAMILY: arial, verdana, helvetica, sans-serif } HTML { FONT-SIZE: 1em; FONT-FAMILY: arial, verdana, helvetica, sans-serif } IMG { FONT-SIZE: 1em; FONT-FAMILY: arial, verdana, helvetica, sans-serif } INPUT { FONT-SIZE: 1em; FONT-FAMILY: arial, verdana, helvetica, sans-serif } SELECT { FONT-SIZE: 1em; FONT-FAMILY: arial, verdana, helvetica, sans-serif } TABLE { FONT-SIZE: 1em; FONT-FAMILY: arial, verdana, helvetica, sans-serif } TD { FONT-SIZE: 1em; FONT-FAMILY: arial, verdana, helvetica, sans-serif } LI { FONT-SIZE: 1em; FONT-FAMILY: arial, verdana, helvetica, sans-serif } .pageText { FONT-SIZE: 1em; FONT-FAMILY: arial, verdana, helvetica, sans-serif } .alertMsg { FONT-WEIGHT: bold; COLOR: #ff0000 } .errMsg { FONT-WEIGHT: bold; COLOR: #ff0000 } .pageHeading { FONT-WEIGHT: bold; FONT-SIZE: 150%; MARGIN: 5px 0px 0px 5px; COLOR: = #8cc919; FONT-FAMILY: arial, verdana, helvetica, sans-serif } .pageHeadingBlack { FONT-WEIGHT: bold; FONT-SIZE: 150%; MARGIN: 5px 0px 0px 5px; COLOR: = black; FONT-FAMILY: arial, helvetica, sans-serif } .pageContent { MARGIN: 5px } #sdLoginBox { PADDING-RIGHT: 0.1em; BACKGROUND-POSITION: left bottom; PADDING-LEFT: = 0.1em; FONT-SIZE: 0.8em; PADDING-BOTTOM: 0.1em; OVERFLOW: hidden; WIDTH: = 29.5em; PADDING-TOP: 0.1em; BACKGROUND-REPEAT: repeat-x; HEIGHT: 6.7em; = BACKGROUND-COLOR: #dddddd } #sdLoginBox INPUT { FONT-SIZE: 0.9em } #sdLoginBox DIV { FONT-SIZE: 1em; FONT-FAMILY: arial, verdana, helvetica, sans-serif } #sdLoginBox HTML { FONT-SIZE: 1em; FONT-FAMILY: arial, verdana, helvetica, sans-serif } #sdLoginBox IMG { FONT-SIZE: 1em; FONT-FAMILY: arial, verdana, helvetica, sans-serif } #sdLoginBox INPUT { FONT-SIZE: 1em; FONT-FAMILY: arial, verdana, helvetica, sans-serif } #sdLoginBox SELECT { FONT-SIZE: 1em; FONT-FAMILY: arial, verdana, helvetica, sans-serif } #sdLoginBox TABLE { FONT-SIZE: 1em; FONT-FAMILY: arial, verdana, helvetica, sans-serif } #sdLoginBox TD { FONT-SIZE: 1em; FONT-FAMILY: arial, verdana, helvetica, sans-serif } #sdLoginBoxFold { OVERFLOW: hidden; TEXT-ALIGN: right } #sdQuickSearch TABLE { FONT-SIZE: 1em; PADDING-BOTTOM: 0.3em } #sdQuickSearch INPUT { FONT-SIZE: 1em } #sdQuickSearch TD { FONT-FAMILY: arial, verdana, helvetica, sans-serif } #sdBody { FONT-SIZE: 75%; PADDING-BOTTOM: 4px; WIDTH: 100%; PADDING-TOP: 4px; = FONT-FAMILY: arial, verdana, helvetica, sans-serif } A.ActionButton { TEXT-DECORATION: none } A.ActionButton:hover { TEXT-DECORATION: underline } A.ActionButton:hover:visited { TEXT-DECORATION: underline } #a:link .Active { TEXT-DECORATION: underline } A:visited .Active { TEXT-DECORATION: underline } .ActionButtonDisabled { COLOR: #999999; TEXT-DECORATION: none } A { TEXT-DECORATION: none } A:link { TEXT-DECORATION: none } A:hover { TEXT-DECORATION: underline } A.linkclosed:hover { COLOR: white; BACKGROUND-COLOR: #777777; TEXT-DECORATION: none } A.linkopen { COLOR: white; BACKGROUND-COLOR: #777777; TEXT-DECORATION: none } A.linkopen:link { COLOR: white; BACKGROUND-COLOR: #777777; TEXT-DECORATION: none } A.linkopen:visited { COLOR: white; BACKGROUND-COLOR: #777777; TEXT-DECORATION: none } A.linkopen:active { COLOR: white; BACKGROUND-COLOR: #777777; TEXT-DECORATION: none } SPAN.linkopen { COLOR: white; BACKGROUND-COLOR: #777777; TEXT-DECORATION: none } .linknoborder IMG { BORDER-RIGHT: medium none; BORDER-TOP: medium none; BORDER-LEFT: medium = none; BORDER-BOTTOM: medium none } .linkdisabled { COLOR: #999999 } INPUT.xpstyle { BORDER-RIGHT: #666666 1px solid; BORDER-TOP: #666666 1px solid; = BORDER-LEFT: #666666 1px solid; BORDER-BOTTOM: #666666 1px solid } INPUT.qsinput { FONT-FAMILY: arial, verdana, helvetica, sans-serif; HEIGHT: 1.6em } TABLE.sdMainNav { BACKGROUND-POSITION: left top; MARGIN-TOP: 0.2em; FONT-SIZE: 1em; = WIDTH: 100%; BORDER-BOTTOM: black 0.1em solid; BACKGROUND-REPEAT: = repeat-x; HEIGHT: 2em } TD.sdMainNav { BACKGROUND-POSITION: left top; BACKGROUND-REPEAT: repeat-x } .sci_dirNav { BACKGROUND-COLOR: #6c9d31 } .sci_dirNavActive { =09 } .blueNav { BACKGROUND-COLOR: #0000aa } .blueNavActive { COLOR: #0000aa; BACKGROUND-COLOR: #bad3f3 } .redNav { BACKGROUND-COLOR: #990000 } .redNavActive { COLOR: #990000; BACKGROUND-COLOR: #f7cece } .greyNav { BACKGROUND-COLOR: #7f7f7f } .greyNavActive { COLOR: #7f7f7f; BACKGROUND-COLOR: #e6e6e6 } .greenNav { BACKGROUND-COLOR: #007703 } .greenNavActive { COLOR: #007703; BACKGROUND-COLOR: #c2edc0 } .orangeNav { BACKGROUND-COLOR: #cc5200 } .orangeNavActive { COLOR: #cc5200; BACKGROUND-COLOR: #ffdfca } TABLE.sdMainNav A:hover { TEXT-DECORATION: none } TABLE.sdMainNavCenter TD { FONT-SIZE: 1em; BORDER-LEFT: white 0.01em solid; HEIGHT: 1.82em } TABLE.sdMainNavCenter TBODY { FONT-SIZE: 1em } TD { FONT-SIZE: 1em } TR { FONT-SIZE: 1em } DIV { FONT-SIZE: 1em } TABLE.sdMainNavCenter { BORDER-RIGHT: white 0.01em solid; HEIGHT: 1.82em } TABLE.sdMainNav A { FONT-WEIGHT: normal; PADDING-BOTTOM: 0.33em; VERTICAL-ALIGN: middle; = COLOR: white; PADDING-TOP: 0.19em; HEIGHT: 1.82em; TEXT-DECORATION: none } TABLE.sdMainNav A:link { FONT-WEIGHT: normal; PADDING-BOTTOM: 0.33em; VERTICAL-ALIGN: middle; = COLOR: white; PADDING-TOP: 0.19em; HEIGHT: 1.82em; TEXT-DECORATION: none } TABLE.sdMainNav A:visited { FONT-WEIGHT: normal; PADDING-BOTTOM: 0.33em; VERTICAL-ALIGN: middle; = COLOR: white; PADDING-TOP: 0.19em; HEIGHT: 1.82em; TEXT-DECORATION: none } DIV.sdMainNavPadding { PADDING-RIGHT: 1em; PADDING-LEFT: 1em } .alphaDivider { DISPLAY: inline; WIDTH: 0px; POSITION: relative; HEIGHT: 0px } .alphaSelect { LEFT: 0px; POSITION: absolute; TOP: -11px; HEIGHT: 14px; = BACKGROUND-COLOR: #777777 } .libBannerText { FONT-SIZE: smaller; FONT-FAMILY: Verdana, Arial, Helvetica, sans-serif } .sci_dirBtn { BACKGROUND-COLOR: #7aa544 } .sci_dirBtnHov { COLOR: #ffffff; BACKGROUND-COLOR: #1d6b2f } .blueBtn { BACKGROUND-COLOR: #0000aa } .blueBtnHov { COLOR: #0000aa; BACKGROUND-COLOR: #bad3f3 } .greenBtn { BACKGROUND-COLOR: #007703 } .greenBtnHov { COLOR: #007703; BACKGROUND-COLOR: #c2edc0 } .greyBtn { BACKGROUND-COLOR: #7f7f7f } .greyBtnHov { COLOR: #7f7f7f; BACKGROUND-COLOR: #e6e6e6 } .redBtn { BACKGROUND-COLOR: #990000 } .redBtnHov { COLOR: #990000; BACKGROUND-COLOR: #f7cece } .orangeBtn { BACKGROUND-COLOR: #cc5200 } .orangeBtnHov { COLOR: #cc5200; BACKGROUND-COLOR: #ffdfca } INPUT.btn { BORDER-RIGHT: #666666 1px solid; PADDING-RIGHT: 1px; BORDER-TOP: = #666666 1px solid; PADDING-LEFT: 1px; FONT-WEIGHT: bold; FONT-SIZE: = 12px; PADDING-BOTTOM: 0px; MARGIN: 0px 1px 0px 0px; VERTICAL-ALIGN: = middle; OVERFLOW: visible; BORDER-LEFT: #666666 1px solid; WIDTH: auto; = CURSOR: pointer; COLOR: #ffffff; PADDING-TOP: 0px; BORDER-BOTTOM: = #666666 1px solid; TEXT-DECORATION: none } INPUT.btnhov { BORDER-RIGHT: #666666 1px solid; PADDING-RIGHT: 1px; BORDER-TOP: = #666666 1px solid; PADDING-LEFT: 1px; FONT-WEIGHT: bold; FONT-SIZE: = 12px; PADDING-BOTTOM: 0px; MARGIN: 0px 1px 0px 0px; VERTICAL-ALIGN: = middle; OVERFLOW: visible; BORDER-LEFT: #666666 1px solid; WIDTH: auto; = CURSOR: pointer; PADDING-TOP: 0px; BORDER-BOTTOM: #666666 1px solid; = TEXT-DECORATION: none } A.articleBtn { BORDER-RIGHT: medium none; PADDING-RIGHT: 20px; BORDER-TOP: medium = none; PADDING-LEFT: 20px; FONT-WEIGHT: bold; FONT-SIZE: 12px; = PADDING-BOTTOM: 1px; MARGIN: 0px; BORDER-LEFT: medium none; CURSOR: = pointer; COLOR: #ffffff; PADDING-TOP: 1px; BORDER-BOTTOM: medium none; = BACKGROUND-COLOR: #999999; TEXT-DECORATION: none } A.articleBtn:hover { BORDER-RIGHT: medium none; PADDING-RIGHT: 20px; BORDER-TOP: medium = none; PADDING-LEFT: 20px; FONT-WEIGHT: bold; FONT-SIZE: 12px; = PADDING-BOTTOM: 1px; MARGIN: 0px; BORDER-LEFT: medium none; CURSOR: = pointer; COLOR: #ffffff; PADDING-TOP: 1px; BORDER-BOTTOM: medium none; = BACKGROUND-COLOR: #999999; TEXT-DECORATION: none } A.articleBtnActive { CURSOR: default; BACKGROUND-COLOR: #7aa544; TEXT-DECORATION: none } A.articleBtnActive:hover { CURSOR: default; BACKGROUND-COLOR: #7aa544; TEXT-DECORATION: none } A.articleBtn:hover { BACKGROUND-COLOR: #7aa544 } A.btn { BORDER-RIGHT: #666666 1px solid; PADDING-RIGHT: 5px; BORDER-TOP: = #666666 1px solid; PADDING-LEFT: 5px; FONT-WEIGHT: bold; FONT-SIZE: = 12px; PADDING-BOTTOM: 1px; MARGIN: 0px 1px 0px 0px; VERTICAL-ALIGN: = middle; BORDER-LEFT: #666666 1px solid; COLOR: #ffffff; PADDING-TOP: = 1px; BORDER-BOTTOM: #666666 1px solid; FONT-FAMILY: arial, verdana, = helvetica, sans-serif; HEIGHT: 1.17em; TEXT-DECORATION: none } A.btn:hover { BORDER-RIGHT: #666666 1px solid; PADDING-RIGHT: 5px; BORDER-TOP: = #666666 1px solid; PADDING-LEFT: 5px; FONT-WEIGHT: bold; FONT-SIZE: = 12px; PADDING-BOTTOM: 1px; MARGIN: 0px 1px 0px 0px; VERTICAL-ALIGN: = middle; BORDER-LEFT: #666666 1px solid; COLOR: #ffffff; PADDING-TOP: = 1px; BORDER-BOTTOM: #666666 1px solid; FONT-FAMILY: arial, verdana, = helvetica, sans-serif; HEIGHT: 1.17em; TEXT-DECORATION: none } A.sci_dirBtn { BACKGROUND-COLOR: #7aa544 } A.sci_dirBtn:hover { COLOR: #ffffff; BACKGROUND-COLOR: #1d6b2f } A.blueBtn { BACKGROUND-COLOR: #0000aa } A.blueBtn:hover { COLOR: #0000aa; BACKGROUND-COLOR: #bad3f3 } A.greenBtn { BACKGROUND-COLOR: #007703 } A.greenBtn:hover { COLOR: #007703; BACKGROUND-COLOR: #c2edc0 } A.greyBtn { BACKGROUND-COLOR: #7f7f7f } A.greyBtn:hover { COLOR: #7f7f7f; BACKGROUND-COLOR: #e6e6e6 } A.redBtn { BACKGROUND-COLOR: #990000 } A.redBtn:hover { COLOR: #990000; 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#007703 2px solid } .greyBorderBtm { BORDER-BOTTOM: #7f7f7f 2px solid } .redBorderBtm { BORDER-BOTTOM: #990000 2px solid } .orangeBorderBtm { BORDER-BOTTOM: #cc5200 2px solid } .tableFmBgGray { FONT-SIZE: 1em; FONT-FAMILY: arial, helvetica, sans-serif; = BACKGROUND-COLOR: #efefef } .tableFmBgGrayNoCell { PADDING-RIGHT: 5px; PADDING-LEFT: 5px; FONT-SIZE: 1em; PADDING-BOTTOM: = 5px; MARGIN: 5px; PADDING-TOP: 5px; FONT-FAMILY: arial, helvetica, = sans-serif; BACKGROUND-COLOR: #efefef } .tableFmBgPad { MARGIN-TOP: 5px; MARGIN-BOTTOM: 5px; PADDING-BOTTOM: 5px; PADDING-TOP: = 5px; BACKGROUND-COLOR: #efefef } .tableFmCell { PADDING-LEFT: 10px; FONT-WEIGHT: bold; PADDING-BOTTOM: 3px; = MARGIN-LEFT: 10px; VERTICAL-ALIGN: top; PADDING-TOP: 3px; TEXT-ALIGN: = right } .tableFmCellTop { FONT-WEIGHT: bold; VERTICAL-ALIGN: top; PADDING-TOP: 0px; TEXT-ALIGN: = right } .tableFmCellAbove { PADDING-RIGHT: 5px; PADDING-LEFT: 5px; FONT-WEIGHT: bold; = PADDING-BOTTOM: 5px; PADDING-TOP: 5px; TEXT-ALIGN: left 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#000000; FONT-FAMILY: arial, = helvetica, sans-serif } A.errMsg { COLOR: #ff0000; TEXT-DECORATION: none } A.errMsg:hover { TEXT-DECORATION: underline } .boldText { FONT-WEIGHT: bold } .sci_dirColor { COLOR: #7aa544 } .blueColor { COLOR: #0000aa } .greenColor { COLOR: #007703 } .greyColor { COLOR: #7f7f7f } .redColor { COLOR: #990000 } .orangeColor { COLOR: #cc5200 } .tablePlainWhite { PADDING-RIGHT: 0px; PADDING-LEFT: 0px; PADDING-BOTTOM: 0px; = PADDING-TOP: 0px; BACKGROUND-COLOR: #ffffff } .inputBox { BORDER-RIGHT: #666666 1px solid; BORDER-TOP: #666666 1px solid; = FONT-SIZE: 1em; VERTICAL-ALIGN: top; BORDER-LEFT: #666666 1px solid; = BORDER-BOTTOM: #666666 1px solid; BACKGROUND-COLOR: #ffffff } A.tabActive { BORDER-RIGHT: medium none; PADDING-RIGHT: 20px; BORDER-TOP: medium = none; PADDING-LEFT: 20px; FONT-WEIGHT: bold; FONT-SIZE: 1em; = PADDING-BOTTOM: 1px; MARGIN: 0px 2px 0px 0px; BORDER-LEFT: medium none; = COLOR: #ffffff; PADDING-TOP: 1px; BORDER-BOTTOM: medium none; = 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