From: Subject: CANCER STEM CELLS: Redefining the Paradigm of Cancer Treatment Strategies -- Dean 6 (3): 140 -- Molecular Interventions Date: Fri, 25 Jan 2008 18:27:43 +0100 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_0000_01C85F7F.F9B604C0" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.3198 This is a multi-part message in MIME format. ------=_NextPart_000_0000_01C85F7F.F9B604C0 Content-Type: text/html; charset="Windows-1252" Content-Transfer-Encoding: quoted-printable Content-Location: http://molinterv.aspetjournals.org/cgi/content/full/6/3/140 CANCER STEM CELLS: Redefining the Paradigm of Cancer = Treatment Strategies -- Dean 6 (3): 140 -- Molecular = Interventions
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Molecular = Interventions=20 6:140-148, (2006)
=A9 American = Society for=20 Pharmacology and Experimental = Therapeutics
10.1124/mi.6.3.5
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3D*=20Cancer
3D*=20Stem=20 = Cells
Review=20

CANCER STEM CELLS: Redefining the Paradigm of Cancer Treatment = Strategies=20

Michael Dean

National Cancer Institute=96Frederick Cancer Research = and=20 Development Center, Bldg 560, Frederick, MD 21702


3D"Formula


3D"    SUMMARY
= TOP
3D"Summary
The Cancer Stem Cell...
Normal and Abnormal Stem...
Stem Cell Activation and...
Research Implications
Treatment Implications
Conclusions
References
 
C= ancer has=20 been known to arise from long-lived cells in the body and to = possess=20 properties in common with undifferentiated, embryonic cells. = Recent=20 findings of a population of cells in solid tumors resembling = stem=20 cells supports a stem cell model of cancer. A scheme in which = all=20 cancers initiate from "activated" stem cells helps bring = together=20 data from genetic, cell biology, and epidemiology studies. = Cancer can=20 arise from embryonic cells in the case of childhood tumors;=20 hormone-activated stem cells in the case of breast cancer; = and=20 following chronic activation of stem cells caused by tissue = damage.=20 This scheme helps explain the failure of many cancer = therapies,=20 points out deficiencies in certain research approaches, and = focuses=20 the problem on a subset of cells that can be explicitly = targeted,=20 leading to more efficient therapy.


3D"    THE CANCER STEM=20 CELL CONCEPT=20
= TOP
= Summary
3D"The Cancer Stem = Cell...
Normal and Abnormal Stem...
Stem Cell Activation and...
Research Implications
Treatment Implications
Conclusions
References
 
T= he idea=20 that cancer cells have properties in common with embryonic = cells is=20 an old one. Cancer cells divide rapidly and are immortal, as = are=20 embryonic stem cells. Embryonal carcinomas and teratocarcinomas = have=20 long fascinated cancer biologists as they are derived from = embryonic=20 cells and give rise to cells of many lineages. Beatrice Mintz = showed=20 in 1975 that teratocarcinomas could give rise to normal = chimeric mice=20 (1= ).=20 Hamburger and Salmon demonstrated that a subpopulation of = cells in a=20 tumor will grow in soft agar, and termed these cells "tumor = stem=20 cells" (2= ).=20 In 1993, Al Knudson proposed that the ability of many tumor=20 suppressor genes to give rise to specific cancers could be = explained=20 by their effects on tissue stem cells (3= ).=20 John Dick proved in 1994 that cancerous stem cells were = present in=20 acute myelogenous leukemia by isolating such cells and = documenting=20 their self-renewing capacity=97the critical property of all = stem cells=20 (4= ).=20 Additionally, alteration in the growth regulation of stem = cells has=20 been invoked to explain the role of the Patched (PTCH) gene = in=20 causing basal cell carcinomas in patients with nevoid basal = cell=20 carcinoma syndrome (5= ).=20

Thus, the term cancer stem cell has arisen to describe cells = that=20 represent a minor population of a tumor and have the property = of=20 self-renewal. Self-renewal involves the replication of a stem = cell=20 generating, in the simplest example, another stem cell and a = cell=20 committed to differentiate. After many cell divisions, a = tumor=20 arising from a single stem cell is composed of cancer stem = cells and=20 non-stem cells that have a limited lifespan. Cancer stem = cells could=20 arise from normal stem cells, from committed cells that = acquire the=20 ability self-renewal capability, or by the fusion of normal = stem=20 cells with cancer cells.

It has been the identification of a small population of cancer=20 stem cells in solid tumors such as breast and brain, however, = that has captivated the field (6= , 7= ).=20 These stem cells appear to be the only cells in the tumor = capable of=20 immortal growth and transplantation of the tumor. Perhaps = more=20 astonishing is the discovery in long established cell lines, = even=20 HeLa cells, of a minor population of cells with stem cell = properties=20 (8= ,=20 9= ).=20 Many researchers now suspect all cancers are composed of a = mixture of=20 renewing stem cells and "committed" cells that continue to=20 proliferate but have a limited life span. The implications of = this=20 concept are important for basic science and well as cancer = therapy.=20 The regrowth of many cancers following therapy could = represent the=20 survival of cancer stem cells, just as the patient=92s hair = follicles=20 and hematopoietic cells recover, because of the survival of=20 hematopoietic and skin stem cells. A fraction of cells in a = tumor are=20 known to survive radiation treatment and cytotoxic drug = exposure=20 (= 10).=20 Stem cells express drug transporters, DNA repair systems, and = are=20 refractory to programmed cell death, all properties that = would serve=20 to allow a cancer cell to resist our efforts to eliminate it = (= 11=96= 13).=20


3D"    NORMAL AND ABNORMAL STEM CELLS
= TOP
= Summary
The Cancer Stem Cell...
3D"Normal and Abnormal = Stem...
Stem Cell Activation and...
Research Implications
Treatment Implications
Conclusions
References
 
T= he=20 earliest cells of an embryo are totipotent stem cells, able = to give=20 rise to all the cell types of the body. These cells = differentiate=20 into more restricted, multipotent stem cells and then to the = further=20 restricted tissue stem cells. All stem cells have the = property of=20 dividing and giving rise to additional stem cells as well as = to cells=20 committed to differentiation that eventually stop dividing or = undergo=20 apoptosis (= 14).=20 Some stem cells, such as hematopoietic stem cells, continue=20 relatively rapid division, to maintain the massive number of=20 lymphocytes and red blood cells needed by the body. Other = stem cells,=20 such as those in the skin and colon, undergo a slow but = constant=20 growth, replenishing the tissue, whereas other tissue stem = cells=20 can remain quiescent to become activated when stimulated by=20 tissue damage or hormone exposure. It becomes obvious then = that=20 the proliferation of all the different types of stem cells in = the adult requires exquisite control and that aberrant = regulation=20 could be hazardous. Cancer stem cells can arise from two = potential=20 pathways: 1) a stem cell losing growth regulation could = directly=20 become a cancer stem cell, or 2) a mature (i.e., = differentiated=20 or committed cell) could acquire the properties of self = renewal=20 and become a cancer stem cell (Figure 1<= IMG=20 height=3D7 alt=3DGo = src=3D"http://molinterv.aspetjournals.org/icons/fig-down.gif"=20 width=3D8 border=3D1>). There is good evidence for both = mechanisms,=20 as well as the possibility that these two pathways might be = utilized=20 in differing proportions at different times within the tumor. = In=20 fact, transfection studies in which human oncogenes transform = murine=20 fibroblasts demonstrate that non-stem cells can be converted = into=20 cancer (stem) cells. An even more outlandish idea is that = normal=20 hematopoietic stem cells may fuse with tissue cells to = generate=20 cancer stem cells (i.e., the formation of an in vivo = hybridoma)=20 (= 15).=20


<= IMG=20 height=3D95 alt=3D"Figure 1" hspace=3D10=20 = src=3D"http://molinterv.aspetjournals.org/content/vol6/issue3/images/smal= l/Fig1_Dean.gif"=20 width=3D200 vspace=3D5 border=3D2>
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  		Figure 1. Two stages = involved in=20 cancer stem cell activation. Cancer requires the = generation of=20 an actively dividing or "activated" stem cell. This cell may = be an=20 actively dividing stem cell, a stem cell stimulated to = divide by=20 tissue damage or inflammation, or from an a differentiated = (mature)=20 cell acquiring the property of self-renewal. Mutations = inactivating=20 tumor suppressor genes (negative regulators of growth) or = activating=20 oncogenes (positive regulators) convert the activated stem = cell into=20 a cancer stem cell. Further mutations are required for tumor = progression and metastasis. =

 = ;

3D"    STEM CELL ACTIVATION AND=20 CANCER
= TOP
= Summary
The Cancer Stem Cell...
Normal and Abnormal Stem...
3D"Stem Cell Activation = and...
Research Implications
Treatment Implications
Conclusions
References
 
A= ctivation=20 of a stem cell can happen in three ways. 1) The stem cell may = be=20 naturally dividing, as in the embryo or the hematopoietic = system; 2)=20 the stem cell can be activated by hormone such as estrogen = for breast=20 and ovarian cells; 3) the stem cell may be stimulated to = divide=20 through tissue damage caused by inflammation, chemical = exposure or=20 infection. With regard to cancer stem cells, it becomes of = central=20 importance for researchers to understand how stem cells take = this=20 first activation step. A model of stem cell activation and = cancer=20 should take into account the many diverse types of cancer and = their=20 varied causes. Such a model in which stem cell "activation" = is the=20 key first step satisfies many observations (Figure 1<= IMG=20 height=3D7 alt=3DGo = src=3D"http://molinterv.aspetjournals.org/icons/fig-up.gif"=20 width=3D8 border=3D1>). Activation could occur through = hormonal=20 stimulation; tissue damage caused by inflammation, radiation, = chemicals, or infections; or inactivation of certain tumor = suppressor=20 genes (Table 1<= IMG=20 height=3D7 alt=3DGo = src=3D"http://molinterv.aspetjournals.org/icons/fig-down.gif"=20 width=3D8 border=3D1>). An activated stem cell would be a = target for=20 further genetic events leading to autonomous growth, loss of = cell=20 cycle regulation, and resistance to apoptosis=97all well = understood=20 properties of cancer cells (= 16).=20 This model accounts for experimental observations on tumor = initiators=20 and promoters (= 17).=20 Most of the environmental agents causing stem-cell activation = would=20 be classified as promoters, non-DNA-reactive agents that = enhance the=20 tumorigenicity of initiators (i.e., mutagens). Some = environmental=20 agents, such as tobacco smoke and UV irradiation, contribute = to both=20 initiation and promotion by stimulating tissue proliferation = and=20 damaging DNA.


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  		Table 1. Routes to = Stem Cell=20 Activation=20
 
Ca= ncers=20 have been proposed to derive from one of three distinct = tissue types:=20 embryonic, conditional growth, or renewal (3= )=20 (Figure 2<= IMG=20 height=3D7 alt=3DGo = src=3D"http://molinterv.aspetjournals.org/icons/fig-down.gif"=20 width=3D8 border=3D1>). Embryonic cancers arise from rapidly = dividing=20 embryonic stem cells; retinoblastoma is the prototype for = this type=20 of cancer (= 18).=20 Retinoblastoma arises in retinoblasts, embryonic cells in the = eye=20 that are highly proliferative. The target cell is a naturally = activated stem cell. The inactivation of the RB1 gene turns = this=20 embryonic stem cell into a cancer stem cell that ignores the = growth=20 regulatory signals that normally arrest the stem cell once = the=20 developing eye is filled with retinoid cells. Other childhood = cancers=20 such as Wilm=92s tumor and leukemias arise from genetic = events in=20 kidney and hematopoietic stem cells, respectively. These = types of=20 cancer require the fewest genetic events, as the target cell = is a=20 fully activated stem cell. Stem cells are also activated by = hormones=20 and/or growth factors in conditional growth tissues, as = described=20 by Knudson (3= ).=20 An example is the breast, which undergoes dramatic growth = during=20 puberty and is then stimulated by estrogen and other hormones = monthly=20 during the woman=92s reproductive life (= 19).=20 Activated stem cells in the breast would be the target cell = for=20 breast cancer. Inactivation of P53 and other genes would = transform=20 breast tissue stem cells into hyperplastic lesions, the = target for=20 further events leading to the progression of pre-malignant = cells to=20 malignancy (Figure 2<= IMG=20 height=3D7 alt=3DGo = src=3D"http://molinterv.aspetjournals.org/icons/fig-down.gif"=20 width=3D8 border=3D1>).


<= IMG=20 height=3D153 alt=3D"Figure 2" hspace=3D10=20 = src=3D"http://molinterv.aspetjournals.org/content/vol6/issue3/images/smal= l/Fig2_Dean.gif"=20 width=3D200 vspace=3D5 border=3D2>
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  		Figure 2. Carcinomas = arising=20 from activated cancer stem cells in different types of = tissue.=20 Embryonic tumors arise from normal embryonic stem cells. = They are=20 already rapidly dividing (activated) and require few = alterations to=20 become malignant. Conditional growth tissues such as the = breast are=20 activated by hormones during puberty and during reproductive = life.=20 Activated stem cells in these tissues suffering tumor = suppressor=20 gene mutations give rise to premalignant lesions progressing = to=20 malignancy. Renewal tissues such as the colon and skin have = dormant=20 or slowly replicating stem cells. Certain tumor suppressor = gene=20 mutations give rise to activated stem cells with = uncontrolled=20 replication. Subsequent mutations give rise to tumor = progression. =

 = ;
Breast=20 cancer is one of the best understood tumors; epidemiological = risk=20 factors, preventive and therapeutic strategies, and genetic = data=20 regarding this disease can be combined into a single scheme = (Figure=20 3<= IMG=20 height=3D7 alt=3DGo = src=3D"http://molinterv.aspetjournals.org/icons/fig-down.gif"=20 width=3D8 border=3D1>). The major breast cancer risk factors involve = hormonal and reproductive events (= 20).=20 Early onset of menarche increases cancer risk, whereas = pregnancies=20 decrease risk. Each of these factors influences either the = number or=20 activation of breast stem cells. The major prevention = strategies=20 currently available similarly affect either estrogen = production or=20 its action on cells. Inhibition of estrogen receptor = expression=20 activation or of estrogen production can prevent breast = cancer in=20 women at high risk. Similarly, oophorectomy also reduces = cancer=20 risk (= 21).=20 BRCA1 and BRCA2 mutations are thought to reduce the = capacity=20 for DNA repair and therefore raise the risk of breast cancer, = by increasing the probability of downstream genetic events = associated=20 with tumor progression. Because BRCA1 and BRCA2 are = not in=20 the main (i.e., estrogen dependent) pathway to tumorigenesis, = they are not frequently mutated in sporadic tumors. =


<= IMG=20 height=3D88 alt=3D"Figure 3" hspace=3D10=20 = src=3D"http://molinterv.aspetjournals.org/content/vol6/issue3/images/smal= l/Fig3_Dean.gif"=20 width=3D200 vspace=3D5 border=3D2>
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  		Figure 3. A = comprehensive model=20 for breast cancer. Applying a stem cell model to breast = cancer=20 helps explain much of the known data. The major risk factors = are=20 hormonal and can act to influence the number and activation = of stem=20 cells. Useful prevention strategies inhibit estrogen = production or=20 action. BRCA1 and BRCA2 germline mutations = increase=20 the likelihood of P53 inactivation (directly or = through=20 inhibition of DNA repair), but the BRCA genes are outside of = the=20 main pathway and thus are not found in most sporadic tumors. =

 = ;
For=20 renewal tissues such as the colon or skin, the stem cells = divide at a=20 very slow rate. In the absence of perturbation of these = tissues,=20 cancer risk is very low. Activation of stem cells in renewal = tissues=20 can occur by inflammation, or tissue damage caused by = radiation,=20 mutagens, and irritants (Figure 2<= IMG=20 height=3D7 alt=3DGo = src=3D"http://molinterv.aspetjournals.org/icons/fig-up.gif"=20 width=3D8 border=3D1>). This chronic tissue damage leads = to an=20 accelerated division of stem cells, increasing the available = number=20 of target cells. In the colon, inflammatory diseases such as = Crohn=20 Disease and inflammatory bowel disease are risk factors for = cancer,=20 as are high fat/low fiber diets (= 22).=20 Patients with the familial adenomatous polyposis coli (FAP) = syndrome=20 have large numbers of colon polyps. Because polyps are = pre-malignant=20 lesions, individuals that have polyps are at very high risk = for colon=20 cancer (= 23).=20 Mutations in APC are responsible for FAP (= 24,=20 = 25)=20 and these patients as well as mice with an equivalent genetic = mutation exhibit increased cellular proliferation in the = colon (= 26=96= 28).=20 The effect of APC mutations might be to increase the=20 proliferation of colonic stem cells (3= ).=20 Consistent with this model, APC is mutated in the vast = majority of sporadic colon tumors (= 29).=20 The later stage events in colon carcinogenesis are well known = and=20 involve mutations in P53, RAS, and other genes (= 29).=20

The major environmental risk factors can be classified by their=20 role in either activating stem cells or contributing to the=20 formation of a pre-malignant lesion (Table 2<= IMG=20 height=3D7 alt=3DGo = src=3D"http://molinterv.aspetjournals.org/icons/fig-down.gif"=20 width=3D8 border=3D1>). Many cancers in solid tissues are = associated=20 with agents causing inflammation or irritation of the cells = in the=20 tissue. Hepatitis B and C viruses are the principal risk = factor for=20 liver cancer, one of the most common cancers in the world = (= 30,=20 = 31).=20 There is no evidence that HBV or HCV contain genes that = directly=20 inactivate tumor suppressor genes; however, patients = persistently=20 infected with these viruses usually suffer from liver = inflammation=20 and extensive cirrhosis. The liver is known to be = particularly=20 adept at regeneration and repairing damaged tissue. Thus,=20 persistent infection leads to chronic inflammation and tissue = damage=20 and continual stimulation of hepatic stem cells.


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  		Table 2. Genetic = Aberrations=20 Involved in Cancers Originating from Stem Cells=20 =
 
As= bestos=20 is a metabolically inert compound remaining in the lungs of=20 individuals exposed to large amounts of the fibers. Asbestos = exposure=20 causes asbestosis, a fibrotic disease of the lung. Asbestos = exposure=20 is also a risk factor for mesothelioma, a cancer of the = mesothelial=20 layer of the lung (= 32).=20 Asbestos also works synergistically with tobacco in = increasing the=20 risk of other lung cancers (= 33).=20 The chronic fibrosis and irritation caused by asbestos could = be=20 expected to continuously stimulate stem cells residing in the = lung=20 and lead to an increased risk of mutational events causing = the=20 formation of a pre-malignant lesion. Similar rationales can = be=20 invoked for the association of infection of the bacterium = H.=20 pylori and metal dust exposure with gastric cancer (= 34,=20 = 35)=20 and the association of chronic pancreatitis and diabetes = mellitus=20 with pancreatic cancer (= 36).=20

Some agents associated with cancer may contribute both to stem=20 cell activation as well as downstream mutations and tumor = suppressor=20 gene inactivation. Human papilloma viruses, particularly = HPV16=20 and 18 are highly associated with cervical cancer (= 37).=20 These viruses contain proteins (i.e., E6 and E7) that inhibit = the p53 and RB1 tumor suppressor proteins (= 38,=20 = 39);=20 however, most women with pathogenic HPV infection do not = develop=20 cervical cancer and so other factors are likely involved. HPV = infection of cervical stem cells leads to the increased=20 proliferation of these cells. Although HPV employs several = strategies=20 to minimize the immune response (= 40)=20 and infected women do not have a visibly inflamed cervix, the = immune=20 system does seem to recognize HPV-infected cells because=20 immunocompromised patients are at increased risk for cervical = cancer=20 (= 41).=20 This chronic proliferation provides an increased pool of = cells=20 susceptible to viral integration and disease = progression.

The associations of smoking and lung cancer and UV exposure = and=20 skin cancer are thought to principally arise from the mutagenic=20 effects of these agents, and these tissues exhibit highly = specific=20 mutations (= 42).=20 Tobacco smoke contains several mutagens and UV irradiation is = clearly=20 mutagenic. Indeed, both long-term tobacco use and long-term = sun=20 exposure are associated with chronic tissue inflammation and = damage.=20 These effects are likely to lead to an increased = proliferation of=20 tissue stem cells as the body tries to repair the damaged = tissue. The=20 constitutively dividing stem cells in these tissues would be = targets=20 for carcinogen-induced mutations, leading to = tumorigenesis.=20


3D"    RESEARCH IMPLICATIONS=20
= TOP
= Summary
The Cancer Stem Cell...
Normal and Abnormal Stem...
Stem Cell Activation and...
3D"Research = Implications
Treatment Implications
Conclusions
References
 
T= he cancer=20 stem cell concept has profound implications for basic cancer=20 research. While cancer biologists have long recognized the=20 heterogeneity of tumor cells, they have assumed that cloned = cells=20 lines are homogeneous. We now know this to be untrue, and the = analysis of tumor cell cultures involves the study of a large = population of committed tumor cells and a small population of = tumor=20 stem cells. Thus, gene expression, microarray, proteomic, and = many=20 phenotypic assays may need to be reinterpreted. Most screens = for=20 cancer cytotoxic drugs have involved cell cultures treated = over short=20 time periods (= 43).=20 A drug specifically inhibiting a tumor stem cell may show = only modest=20 activity in such an assay and not be chosen for follow up = study.=20 Clinical studies of stem cell inhibitors may need to be = redesigned to=20 capture long term benefits.

Through the use of specific cell surface markers tumor stem = cells=20 might be isolated from cell lines and characterized. Alternatively=20 normal stem cells have been purified by taking advantage of = the=20 fact that stem cells express transport proteins that efflux = certain=20 dyes such as Hoechst and rhodamine. These so called = side-population=20 (SP) cells have been found in bone marrow and normal tissues, = as well=20 as in tumor cell lines (8= , 9= ).=20 The isolation of such tumor stem cells would likely lead to = improved=20 understanding of gene expression, cell surface proteins, and = drug=20 sensitivities of these cells. One long-recognized property of = cancer=20 cells is anchorage independence=97as measured in vitro by = the=20 ability of tumor cells to proliferate into colonies in soft=20 agar medium (2= ).=20 Interestingly, the only normal cells with this property are = stem=20 cells, and only a fraction of cells in a tumor cell culture = can form=20 a colony in soft agar. This clonogenicity ranges from less = that 1% to=20 20%=9650% (Michael Alley, personal communication). A likely = explanation=20 of these data is that the only cells in a tumor cell line = forming=20 colonies are the tumor stem cells, the rest of the cells in = the tumor=20 being committed to differentiation or senescence. In fact, = the=20 fraction of (SP) cells in tumor cell lines correlates with = their=20 clonogenicty, in the few lines that have been examined [data = not=20 shown but reported in (9= )].=20 An exciting implication of this result is that by screening = for drugs=20 suppressing colony formation, we can identify drugs that = inhibit the=20 proliferation of tumor stem cells. Such an assay would be = more=20 cumbersome and have a lower throughput than screens on = adherent cells=20 but could prove to be more informative.

Stem cells, especially germ cells, possess the ability to = migrate=20 throughout the developing body. This is particularly true of=20 cells derived from the neural crest that give rise to = melanoma,=20 neuroblastoma, and small cell lung cancers. Most of the = mortality=20 of cancer arises from the metastasis of cells from the = original=20 tumor site, and their migration to other organs. Could the = metastatic=20 ability of some tumor cells relate to an innate property of=20 stem cells to migrate? If so, then further understanding of=20 germ cell migration could lead to new insights into = metastasis.=20


3D"    TREATMENT IMPLICATIONS=20
= TOP
= Summary
The Cancer Stem Cell...
Normal and Abnormal Stem...
Stem Cell Activation and...
Research Implications
3D"Treatment = Implications
Conclusions
References
 
I= n the=20 light of a stem cell model, tumors that regress following = radiation=20 and chemotherapy protocols are not the result of a rare cell = from=20 within the tumor but are the regrowth of the cancer stem = cells in the=20 original tumor. However, tumor stem cells could acquire = additional=20 genetic changes to render them even more drug resistant, = radiation=20 resistant, or aneuploid. Because cures are achieved for many = types of=20 cancer, cancer stem cells must be eliminated by some = therapeutic=20 strategies. The committed cells in a tumor may very well play = a role=20 in supporting or stimulating the stem cells, and regression = of=20 the committed cells in a tumor could lead to elimination of = the=20 stem cells as well. Immune surveillance is clearly important = in many=20 cancers (= 44),=20 and decreasing the mass of the tumor may allow the immune = system to=20 efficiently recognize the remaining cells. Clearly, targeted=20 therapies that directly suppress or kill the tumor stem cells = may=20 have increased efficacy. For example, angiogenesis is likely = to be=20 critical to provide blood supply to the tumor stem cells, so=20 strategies to fight the development or inhibit the tumors = blood=20 vessels are likely to be effective (= 45).=20

Therapies that target cancer stem cells, however, may require = new=20 approaches to evaluating the success of a cancer agent. If a = drug=20 only inhibits the proliferation of the cancer stem cells = (rather than=20 killing them outright) the drug will affect the growth of = only a=20 minor population of cells and there may be a minimal decrease = in the=20 growth of the tumor in the short term. Pharmacodynamic assays = to=20 demonstrate the drug is inhibiting the target protein will be = important and the use of endpoints such as disease free = survival may=20 be useful. Cancer stem cell=96targeted therapies will no = doubt be=20 combined with standard therapies, to attempt the complete = elimination=20 of the tumor. As with all cancer therapies it will be = critical to=20 establish a dose and schedule where the tumor is suppressed = or=20 eliminated without undue toxicity of normal stem cells. = Recent data=20 on mouse leukemia models suggest that such therapies may be=20 established (= 46).=20

One of the innate resistance mechanisms of stem cells is the=20 expression of one or more ATP-binding cassette (ABC) = transporters.=20 These pumps play a role in protecting stem cells from = xenobiotic=20 toxins (= 47).=20 ABCG2 and ABCB1/MDR1 genes are expressed in the = vast=20 majority of stem cells and in most tumor stem cells (= 11=96= 13).=20 These transporters can efflux fluorescent dyes such as = rhodamine=20 and Hoecht 33342, and this property allows the stem cells to = be=20 separated from non-stem cells on a cell sorter (= 48).=20 The combined use of chemotherapy drugs and ABC transporter=20 inhibitors could be used to specifically target cancer stem = cells=20 (= 49,=20 = 50).=20 In fact, there are highly specific inhibitors of ABCB1 in = clinical=20 use and ABCG2 inhibitors in development (= 51).=20 These therapies would be predicted to have toxic effects on = the=20 patient=92s normal stem cells, and both ABCG2 and ABCB1 play = a role in=20 the blood-brain barrier, which suggests that this approach = would=20 have to be carefully titrated to avoid excessive toxicity. =

Directly targeting the growth of stem cells could be a fruitful=20 avenue. Because of ground-breaking work in Drosophila and=20 other developmental systems, we know a great deal about the=20 growth regulatory pathways operative in embryonic cells = (= 52).=20 One such pathway, involving the Hedgehog (HH) and WNT = signaling=20 molecules, contains a large number of genes that can act as = tumor=20 suppressor genes or oncogenes in mammalian cells (5= ).=20 For example, Patched (PTCH) codes for the receptor = that binds=20 HH molecules and is mutated in patients with nevoid basal = cell=20 carcinoma syndrome (= 53,=20 = 54).=20 PTCH is also mutated in nearly all sporadic basal cell = carcinomas=20 and in some medulloblastomas (= 55).=20 The mammalian HH genes (sonic hedgehog (SHH), Indian hedgehog = (IHH),=20 and desert hedgehog (DHH)) are overexpressed in a wide = variety of=20 cancers, including small-cell lung, pancreas, gastric, = breast, and=20 prostate (= 56=96= 59).=20 HH family overexpression and PTCH mutation both have the = effect=20 of constitutive action of smoothened (SMO) a G = protein=96coupled=20 receptor that is a key signaling component of the pathway.=20 Constitutive HH family expression could lead to stem = cell=20 activation and appears to be a common feature of many = cancers.=20

Cyclopamine is a compound originally discovered in the Corn = Lily=20 (Veratrum californicum), a plant teratogenic to sheep (= 60).=20 Cyclopamine specifically inhibits the SMO protein and can = suppress=20 the growth of cells and tumors that contain activated HH (= 61).=20 Human prostate tumors grown as xenografts in mice were = completely=20 eliminated following twenty-one days of treatment with = cyclopamine=20 (= 59),=20 and UV-induced basal cell carcinomas were suppressed in mice = given=20 low levels of cyclopamine in their drinking water (= 62).=20 Both cyclopamine and SMO inhibitors are in development as = anti-cancer=20 agents.

The Notch protein is another protein important to the growth = and=20 differentiation of stem cells. Notch is processed by the = enzyme -secretase, the same enzyme that processes the APP = protein=20 important in Alzheimer Disease. Several companies have = developed=20 3D{gamma}-secretase inhibitors and these drugs may have = applications=20 to cancer therapy. Rapamycin is a bacterially derived = molecule=20 that inhibits the target of rapamycin (TOR) family of kinases = (= 63).=20 Rapamycin has immunosuppressive properties and is also used = to treat=20 certain leukemias (= 64).=20 Yilmaz et al. have shown that rapamycin can selectively = inhibit=20 leukemia initiating cells in leukemias generated in mice = harboring a=20 conditional deletion of PTEN (phosphatase and tensin homolog = deleted=20 on chromosome 10), a phosphati-dylinositol phosphate (PIP)=20 phosphatase often mutated in human tumors (= 46).=20 Rapamycin depletes the leukemia-initiating cells while = restoring the=20 normal hematopoieitc stem cells, suggesting therapies can be = devised=20 specific for cancer stem cells. Additional strategies to = directly=20 target the growth of tumor stem cells can now be developed = and may=20 prove superior in effectiveness.

Nearly all cancer mortality derives from metastatic leasions, = not=20 the primary tumor. Therefore it is of great importance to = understand=20 the nature of metastases and how to inhibit or eliminate = them. It=20 follows from the cancer stem cell model that metastatic = lesions must=20 initiate from a cancer stem cell. In fact many stem cells, = such as=20 those in the neural crest, migrate through the developing = body. Thus=20 the migratory behavior that we recognize as metastasis may be = a=20 "normal" function of the cancer stem cell. Therefore drugs = that=20 inhibit the growth of cancer stem cells may have the effect = of=20 suppressing metastasis. Although the ultimate goal of cancer = therapy=20 is the complete elimination of the tumor, turning cancer from = an=20 acute disease to a chromic disease may be a more achievable = near-term=20 prospect.


3D"    CONCLUSIONS
= TOP
= Summary
The Cancer Stem Cell...
Normal and Abnormal Stem...
Stem Cell Activation and...
Research Implications
Treatment Implications
3D"Conclusions
References
 
T= he=20 concept of cancer as a stem cell disease has the potential to = dramatically change our view of the problem. By separating = the=20 disease into a stem cell activation phase and a tumor progression=20 phase, historical cancer studies can be reinterpreted with = new=20 understanding. Research efforts to understand the growth of=20 tumor stem cells as well as identify tumor stem cell antigens = could lead to new targeted approaches. Cancer diagnostics,=20 prevention, and therapeutics are likely to be greatly aided = by this=20 new insight. Normal stem cells are being intensively studied = to=20 develop approaches for replacing damaged cells and tissues in = the body. The insight from this work is likely to aid the=20 understanding of cancer stem cells.


Michael=20 Dean, PhD, received his doctorate degree from = the=20 Biochemistry Department at the Boston University = School=20 of Medicine. His advisor was Dr. Gail Sonenshein = and he=20 studied transcription and regulation of the MYC = oncogene.=20 He performed his postdoctoral studies at the=20 NCI-Frederick with Dr. George Vande Woude where he = studied=20 the MET oncogene. They discovered linkage of MET to = the=20 locus for cystic fibrosis. After joining the = Laboratory=20 of Genomic Diversity (LGD), Dr. Dean=92s group = participated=20 in the molecular cloning of the cystic fibrosis = gene,=20 identification of variants in CCR5 and CCR2 = involved in=20 HIV-1 infection and AIDS progression, = identification of=20 the VHL and PTCH genes in inherited cancers, and = the=20 involvement of several ATP-binding cassette (ABC) = transporters in human disease and drug resistance. = He is=20 shown here teaching as part of the Elementary Outreach=20 Program. This is an effort Michael created bringing = NCI=20 and contract employees into Frederick County = elementary=20 schools. Over 7,000 students have received = hands-on=20 science lessons in the six years the program has = been=20 operating. In his spare time Michael enjoys = kayaking,=20 skiing and playing softball. He is Chief of the = Human=20 Genetics Section in the LGD. E-mail: dean{at}ncifcrf.gov ; fax: 301-846-1909. =


3D"   =20 References
= TOP
= Summary
The Cancer Stem Cell...
Normal and Abnormal Stem...
Stem Cell Activation and...
Research Implications
Treatment Implications
Conclusions
3D"References
&nb= sp;

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Opera = requires=0A= the Sun Java Runtime Environment .=0A= =0A= by Andrew Gregory=0A= http://www.scss.com.au/family/andrew/webdesign/xmlhttprequest/=0A= =0A= This work is licensed under the Creative Commons Attribution License. To = view a=0A= copy of this license, visit = http://creativecommons.org/licenses/by-sa/2.5/ or=0A= send a letter to Creative Commons, 559 Nathan Abbott Way, Stanford, = California=0A= 94305, USA.=0A= =0A= Attribution: Leave my name and web address in this script intact.=0A= =0A= Not Supported in Opera=0A= ----------------------=0A= * user/password authentication=0A= * responseXML data member=0A= =0A= Not Fully Supported in Opera=0A= ----------------------------=0A= * async requests=0A= * abort()=0A= * getAllResponseHeaders(), getAllResponseHeader(header)=0A= =0A= */=0A= // IE support=0A= if (window.ActiveXObject && !window.XMLHttpRequest) {=0A= window.XMLHttpRequest =3D function() {=0A= var msxmls =3D new Array(=0A= 'Msxml2.XMLHTTP.5.0',=0A= 'Msxml2.XMLHTTP.4.0',=0A= 'Msxml2.XMLHTTP.3.0',=0A= 'Msxml2.XMLHTTP',=0A= 'Microsoft.XMLHTTP');=0A= for (var i =3D 0; i < msxmls.length; i++) {=0A= try {=0A= return new ActiveXObject(msxmls[i]);=0A= } catch (e) {=0A= }=0A= }=0A= return null;=0A= };=0A= }=0A= // Gecko support=0A= /* ;-) */=0A= // Opera support=0A= if (window.opera && !window.XMLHttpRequest) {=0A= window.XMLHttpRequest =3D function() {=0A= this.readyState =3D 0; // = 0=3Duninitialized,1=3Dloading,2=3Dloaded,3=3Dinteractive,4=3Dcomplete=0A= this.status =3D 0; // HTTP status codes=0A= this.statusText =3D '';=0A= this._headers =3D [];=0A= this._aborted =3D false;=0A= this._async =3D true;=0A= this._defaultCharset =3D 'ISO-8859-1';=0A= this._getCharset =3D function() {=0A= var charset =3D _defaultCharset;=0A= var contentType =3D = this.getResponseHeader('Content-type').toUpperCase();=0A= val =3D contentType.indexOf('CHARSET=3D');=0A= if (val !=3D -1) {=0A= charset =3D contentType.substring(val);=0A= }=0A= val =3D charset.indexOf(';');=0A= if (val !=3D -1) {=0A= charset =3D charset.substring(0, val);=0A= }=0A= val =3D charset.indexOf(',');=0A= if (val !=3D -1) {=0A= charset =3D charset.substring(0, val);=0A= }=0A= return charset;=0A= };=0A= this.abort =3D function() {=0A= this._aborted =3D true;=0A= };=0A= this.getAllResponseHeaders =3D function() {=0A= return this.getAllResponseHeader('*');=0A= };=0A= this.getAllResponseHeader =3D function(header) {=0A= var ret =3D '';=0A= for (var i =3D 0; i < this._headers.length; i++) {=0A= if (header =3D=3D '*' || this._headers[i].h =3D=3D header) {=0A= ret +=3D this._headers[i].h + ': ' + this._headers[i].v + '\n';=0A= }=0A= }=0A= return ret;=0A= };=0A= this.getResponseHeader =3D function(header) {=0A= var ret =3D getAllResponseHeader(header);=0A= var i =3D ret.indexOf('\n');=0A= if (i !=3D -1) {=0A= ret =3D ret.substring(0, i);=0A= }=0A= return ret;=0A= };=0A= this.setRequestHeader =3D function(header, value) {=0A= this._headers[this._headers.length] =3D {h:header, v:value};=0A= };=0A= this.open =3D function(method, url, async, user, password) {=0A= this.method =3D method;=0A= this.url =3D url;=0A= this._async =3D true;=0A= this._aborted =3D false;=0A= this._headers =3D [];=0A= if (arguments.length >=3D 3) {=0A= this._async =3D async;=0A= }=0A= if (arguments.length > 3) {=0A= opera.postError('XMLHttpRequest.open() - user/password not = supported');=0A= }=0A= this.readyState =3D 1;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= };=0A= this.send =3D function(data) {=0A= if (!navigator.javaEnabled()) {=0A= alert("XMLHttpRequest.send() - Java must be installed and = enabled.");=0A= return;=0A= }=0A= if (this._async) {=0A= setTimeout(this._sendasync, 0, this, data);=0A= // this is not really asynchronous and won't execute until the = current=0A= // execution context ends=0A= } else {=0A= this._sendsync(data);=0A= }=0A= }=0A= this._sendasync =3D function(req, data) {=0A= if (!req._aborted) {=0A= req._sendsync(data);=0A= }=0A= };=0A= this._sendsync =3D function(data) {=0A= this.readyState =3D 2;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= // open connection=0A= var url =3D new java.net.URL(new = java.net.URL(window.location.href), this.url);=0A= var conn =3D url.openConnection();=0A= for (var i =3D 0; i < this._headers.length; i++) {=0A= conn.setRequestProperty(this._headers[i].h, this._headers[i].v);=0A= }=0A= this._headers =3D [];=0A= if (this.method =3D=3D 'POST') {=0A= // POST data=0A= conn.setDoOutput(true);=0A= var wr =3D new = java.io.OutputStreamWriter(conn.getOutputStream(), this._getCharset());=0A= wr.write(data);=0A= wr.flush();=0A= wr.close();=0A= }=0A= // read response headers=0A= // NOTE: the getHeaderField() methods always return nulls for me :(=0A= var gotContentEncoding =3D false;=0A= var gotContentLength =3D false;=0A= var gotContentType =3D false;=0A= var gotDate =3D false;=0A= var gotExpiration =3D false;=0A= var gotLastModified =3D false;=0A= for (var i =3D 0; ; i++) {=0A= var hdrName =3D conn.getHeaderFieldKey(i);=0A= var hdrValue =3D conn.getHeaderField(i);=0A= if (hdrName =3D=3D null && hdrValue =3D=3D null) {=0A= break;=0A= }=0A= if (hdrName !=3D null) {=0A= this._headers[this._headers.length] =3D {h:hdrName, = v:hdrValue};=0A= switch (hdrName.toLowerCase()) {=0A= case 'content-encoding': gotContentEncoding =3D true; break;=0A= case 'content-length' : gotContentLength =3D true; break;=0A= case 'content-type' : gotContentType =3D true; break;=0A= case 'date' : gotDate =3D true; break;=0A= case 'expires' : gotExpiration =3D true; break;=0A= case 'last-modified' : gotLastModified =3D true; break;=0A= }=0A= }=0A= }=0A= // try to fill in any missing header information=0A= var val;=0A= val =3D conn.getContentEncoding();=0A= if (val !=3D null && !gotContentEncoding) = this._headers[this._headers.length] =3D {h:'Content-encoding', v:val};=0A= val =3D conn.getContentLength();=0A= if (val !=3D -1 && !gotContentLength) = this._headers[this._headers.length] =3D {h:'Content-length', v:val};=0A= val =3D conn.getContentType();=0A= if (val !=3D null && !gotContentType) = this._headers[this._headers.length] =3D {h:'Content-type', v:val};=0A= val =3D conn.getDate();=0A= if (val !=3D 0 && !gotDate) this._headers[this._headers.length] = =3D {h:'Date', v:(new Date(val)).toUTCString()};=0A= val =3D conn.getExpiration();=0A= if (val !=3D 0 && !gotExpiration) = this._headers[this._headers.length] =3D {h:'Expires', v:(new = Date(val)).toUTCString()};=0A= val =3D conn.getLastModified();=0A= if (val !=3D 0 && !gotLastModified) = this._headers[this._headers.length] =3D {h:'Last-modified', v:(new = Date(val)).toUTCString()};=0A= // read response data=0A= var reqdata =3D '';=0A= var stream =3D conn.getInputStream();=0A= if (stream) {=0A= var reader =3D new java.io.BufferedReader(new = java.io.InputStreamReader(stream, this._getCharset()));=0A= var line;=0A= while ((line =3D reader.readLine()) !=3D null) {=0A= if (this.readyState =3D=3D 2) {=0A= this.readyState =3D 3;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= }=0A= reqdata +=3D line + '\n';=0A= }=0A= reader.close();=0A= this.status =3D 200;=0A= this.statusText =3D 'OK';=0A= this.responseText =3D reqdata;=0A= this.readyState =3D 4;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= if (this.onload) {=0A= this.onload();=0A= }=0A= } else {=0A= // error=0A= this.status =3D 404;=0A= this.statusText =3D 'Not Found';=0A= this.responseText =3D '';=0A= this.readyState =3D 4;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= if (this.onerror) {=0A= this.onerror();=0A= }=0A= }=0A= };=0A= };=0A= }=0A= // ActiveXObject emulation=0A= if (!window.ActiveXObject && window.XMLHttpRequest) {=0A= window.ActiveXObject =3D function(type) {=0A= switch (type.toLowerCase()) {=0A= case 'microsoft.xmlhttp':=0A= case 'msxml2.xmlhttp':=0A= case 'msxml2.xmlhttp.3.0':=0A= case 'msxml2.xmlhttp.4.0':=0A= case 'msxml2.xmlhttp.5.0':=0A= return new XMLHttpRequest();=0A= }=0A= return null;=0A= };=0A= }=0A= ------=_NextPart_000_0000_01C85F7F.F9B604C0 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://molinterv.aspetjournals.org/javascript/ajax/utility.js /************************************************************************= *****=0A= * javascript/ajax/utility.js=0A= *=0A= * Utility functions for working with XMLHttpRequest data.=0A= *=0A= * Copyright 2006 Board of Trustees of the Leland Stanford Junior = University.=0A= = *************************************************************************= ***/=0A= =0A= /*=0A= * Copy XML nodes into an HTMLElement. This effectively=0A= * clones XML markup which uses XHTML naming conventions=0A= * into an HTML DOM.=0A= */=0A= function copy_xml_to_html(src, dst) {=0A= if (src.nodeType =3D=3D 1) { /* Node.ELEMENT_NODE */=0A= var e =3D document.createElement(src.nodeName);=0A= for (var i =3D 0; i < src.childNodes.length; i++) {=0A= copy_xml_to_html(src.childNodes[i], e);=0A= }=0A= for (var i =3D 0; i < src.attributes.length; i++) {=0A= var n =3D src.attributes[i].name;=0A= var v =3D unescape_xml_string(src.attributes[i].value); =0A= e.setAttribute(n, v);=0A= if (n =3D=3D "class") {=0A= e.className =3D v;=0A= }=0A= else if (n =3D=3D "style") {=0A= set_css_style(v, e, "");=0A= }=0A= }=0A= dst.appendChild(e);=0A= }=0A= else if (src.nodeType =3D=3D 3) { /* Node.TEXT_NODE */=0A= dst.appendChild(document.createTextNode(src.nodeValue));=0A= }=0A= }=0A= =0A= /* =0A= * It is unclear that this is the right thing to be calling=0A= * from copy_xml_to_html, but it appears that Safari decides=0A= * to convert & to the NCR #, and then encodes that=0A= * NCR to &%26%2338;. So, I'm going to treat the DOM Attr=0A= * value as a plain string, and run our XML string input=0A= * through the decoding routine below.=0A= */=0A= function unescape_xml_string(s) {=0A= return s.replace(/'/g, "'")=0A= .replace(/'/g, "'")=0A= .replace(/"/g, "\"")=0A= .replace(/"/g, "\"")=0A= .replace(/>/g, ">")=0A= .replace(/>/g, ">")=0A= .replace(/</g, "<")=0A= .replace(/</g, "<")=0A= .replace(/&/g, "&")=0A= .replace(/&/g, "&");=0A= }=0A= =0A= /*=0A= * Parse set of CSS rules and apply them to an element.=0A= * This is quite horrifying, but I'm unable to determine=0A= * how else to handle this with IE 6. FireFox and other=0A= * sane browsers let you simply set the style attribute=0A= * or use e.style.setProperty(rule, value, priority),=0A= * IE 6 appears to have neither of these capabilities..=0A= */=0A= function set_css_style(css, e, priority) {=0A= var rules =3D css.split(";");=0A= for (var i =3D 0; i < rules.length; i++) {=0A= var nvpair =3D rules[i].split(":");=0A= if (nvpair.length =3D=3D 2) {=0A= try {=0A= var name =3D nvpair[0]; /* style attribute */=0A= var value =3D nvpair[1]; /* attribute value */=0A= =0A= /*=0A= * For each possible style attribute, set the=0A= * appropriate style property in the element.=0A= */=0A= if (name =3D=3D "background") {=0A= e.style.background =3D value;=0A= }=0A= else if (name =3D=3D "background-attachment") {=0A= e.style.backgroundAttachment =3D value;=0A= }=0A= else if (name =3D=3D "background-color") {=0A= e.style.backgroundColor =3D value;=0A= }=0A= else if (name =3D=3D "background-image") {=0A= e.style.backgroundImage =3D value;=0A= }=0A= else if (name =3D=3D "background-position") {=0A= e.style.backgroundPosition =3D value;=0A= }=0A= else if (name =3D=3D "background-position-x") {=0A= e.style.backgroundPositionX =3D value;=0A= }=0A= else if (name =3D=3D "background-position-y") {=0A= e.style.backgroundPositionY =3D value;=0A= }=0A= else if (name =3D=3D "background-repeat") {=0A= e.style.backgroundRepeat =3D value;=0A= }=0A= else if (name =3D=3D "behavior") {=0A= e.style.behavior =3D value;=0A= }=0A= else if (name =3D=3D "border") {=0A= e.style.border =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom") {=0A= e.style.borderBottom =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom-color") {=0A= e.style.borderBottomColor =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom-style") {=0A= e.style.borderBottomStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom-width") {=0A= e.style.borderBottomWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-collapse") {=0A= e.style.borderCollapse =3D value;=0A= }=0A= else if (name =3D=3D "border-color") {=0A= e.style.borderColor =3D value;=0A= }=0A= else if (name =3D=3D "border-left") {=0A= e.style.borderLeft =3D value;=0A= }=0A= else if (name =3D=3D "border-left-color") {=0A= e.style.borderLeftColor =3D value;=0A= }=0A= else if (name =3D=3D "border-left-style") {=0A= e.style.borderLeftStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-left-width") {=0A= e.style.borderLeftWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-right") {=0A= e.style.borderRight =3D value;=0A= }=0A= else if (name =3D=3D "border-right-color") {=0A= e.style.borderRightColor =3D value;=0A= }=0A= else if (name =3D=3D "border-right-style") {=0A= e.style.borderRightStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-right-width") {=0A= e.style.borderRightWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-style") {=0A= e.style.borderStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-top") {=0A= e.style.borderTop =3D value;=0A= }=0A= else if (name =3D=3D "border-top-color") {=0A= e.style.borderTopColor =3D value;=0A= }=0A= else if (name =3D=3D "border-top-style") {=0A= e.style.borderTopStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-top-width") {=0A= e.style.borderTopWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-width") {=0A= e.style.borderWidth =3D value;=0A= }=0A= else if (name =3D=3D "bottom") {=0A= e.style.bottom =3D value;=0A= }=0A= else if (name =3D=3D "clear") {=0A= e.style.clear =3D value;=0A= }=0A= else if (name =3D=3D "clip") {=0A= e.style.clip =3D value;=0A= }=0A= else if (name =3D=3D "color") {=0A= e.style.color =3D value;=0A= }=0A= else if (name =3D=3D "cssText") {=0A= e.style.Sets =3D value;=0A= }=0A= else if (name =3D=3D "cursor") {=0A= e.style.cursor =3D value;=0A= }=0A= else if (name =3D=3D "direction") {=0A= e.style.direction =3D value;=0A= }=0A= else if (name =3D=3D "display") {=0A= e.style.display =3D value;=0A= }=0A= else if (name =3D=3D "font") {=0A= e.style.font =3D value;=0A= }=0A= else if (name =3D=3D "font-family") {=0A= e.style.fontFamily =3D value;=0A= }=0A= else if (name =3D=3D "font-size") {=0A= e.style.fontSize =3D value;=0A= }=0A= else if (name =3D=3D "font-style") {=0A= e.style.fontStyle =3D value;=0A= }=0A= else if (name =3D=3D "font-variant") {=0A= e.style.fontVariant =3D value;=0A= }=0A= else if (name =3D=3D "font-weight") {=0A= e.style.fontWeight =3D value;=0A= }=0A= else if (name =3D=3D "height") {=0A= e.style.height =3D value;=0A= }=0A= else if (name =3D=3D "ime-mode") {=0A= e.style.imeMode =3D value;=0A= }=0A= else if (name =3D=3D "layout-flow") {=0A= e.style.layoutFlow =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid") {=0A= e.style.layoutGrid =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-char") {=0A= e.style.layoutGridChar =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-line") {=0A= e.style.layoutGridLine =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-mode") {=0A= e.style.layoutGridMode =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-type") {=0A= e.style.layoutGridType =3D value;=0A= }=0A= else if (name =3D=3D "left") {=0A= e.style.left =3D value;=0A= }=0A= else if (name =3D=3D "letter-spacing") {=0A= e.style.letterSpacing =3D value;=0A= }=0A= else if (name =3D=3D "line-break") {=0A= e.style.lineBreak =3D value;=0A= }=0A= else if (name =3D=3D "line-height") {=0A= e.style.lineHeight =3D value;=0A= }=0A= else if (name =3D=3D "list-style") {=0A= e.style.listStyle =3D value;=0A= }=0A= else if (name =3D=3D "list-style-image") {=0A= e.style.listStyleImage =3D value;=0A= }=0A= else if (name =3D=3D "list-style-position") {=0A= e.style.listStylePosition =3D value;=0A= }=0A= else if (name =3D=3D "list-style-type") {=0A= e.style.listStyleType =3D value;=0A= }=0A= else if (name =3D=3D "margin") {=0A= e.style.margin =3D value;=0A= }=0A= else if (name =3D=3D "margin-bottom") {=0A= e.style.marginBottom =3D value;=0A= }=0A= else if (name =3D=3D "margin-left") {=0A= e.style.marginLeft =3D value;=0A= }=0A= else if (name =3D=3D "margin-right") {=0A= e.style.marginRight =3D value;=0A= }=0A= else if (name =3D=3D "margin-top") {=0A= e.style.marginTop =3D value;=0A= }=0A= else if (name =3D=3D "min-height") {=0A= e.style.minHeight =3D value;=0A= }=0A= else if (name =3D=3D "overflow") {=0A= e.style.overflow =3D value;=0A= }=0A= else if (name =3D=3D "overflow-x") {=0A= e.style.overflowX =3D value;=0A= }=0A= else if (name =3D=3D "overflow-y") {=0A= e.style.overflowY =3D value;=0A= }=0A= else if (name =3D=3D "padding") {=0A= e.style.padding =3D value;=0A= }=0A= else if (name =3D=3D "padding-bottom") {=0A= e.style.paddingBottom =3D value;=0A= }=0A= else if (name =3D=3D "padding-left") {=0A= e.style.paddingLeft =3D value;=0A= }=0A= else if (name =3D=3D "padding-right") {=0A= e.style.paddingRight =3D value;=0A= }=0A= else if (name =3D=3D "padding-top") {=0A= e.style.paddingTop =3D value;=0A= }=0A= else if (name =3D=3D "page-break-after") {=0A= e.style.pageBreakAfter =3D value;=0A= }=0A= else if (name =3D=3D "page-break-before") {=0A= e.style.pageBreakBefore =3D value;=0A= }=0A= else if (name =3D=3D "pixelBottom") {=0A= e.style.pixelBottom =3D value;=0A= }=0A= else if (name =3D=3D "pixelHeight") {=0A= e.style.pixelHeight =3D value;=0A= }=0A= else if (name =3D=3D "pixelLeft") {=0A= e.style.pixelLeft =3D value;=0A= }=0A= else if (name =3D=3D "pixelRight") {=0A= e.style.pixelRight =3D value;=0A= }=0A= else if (name =3D=3D "pixelTop") {=0A= e.style.pixelTop =3D value;=0A= }=0A= else if (name =3D=3D "pixelWidth") {=0A= e.style.pixelWidth =3D value;=0A= }=0A= else if (name =3D=3D "posBottom") {=0A= e.style.posBottom =3D value;=0A= }=0A= else if (name =3D=3D "posHeight") {=0A= e.style.posHeight =3D value;=0A= }=0A= else if (name =3D=3D "position") {=0A= e.style.position =3D value;=0A= }=0A= else if (name =3D=3D "posLeft") {=0A= e.style.posLeft =3D value;=0A= }=0A= else if (name =3D=3D "posRight") {=0A= e.style.posRight =3D value;=0A= }=0A= else if (name =3D=3D "posTop") {=0A= e.style.posTop =3D value;=0A= }=0A= else if (name =3D=3D "posWidth") {=0A= e.style.posWidth =3D value;=0A= }=0A= else if (name =3D=3D "right") {=0A= e.style.right =3D value;=0A= }=0A= else if (name =3D=3D "ruby-align") {=0A= e.style.rubyAlign =3D value;=0A= }=0A= else if (name =3D=3D "ruby-overhang") {=0A= e.style.rubyOverhang =3D value;=0A= }=0A= else if (name =3D=3D "ruby-position") {=0A= e.style.rubyPosition =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-3dlight-color") {=0A= e.style.scrollbar3dLightColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-arrow-color") {=0A= e.style.scrollbarArrowColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-base-color") {=0A= e.style.scrollbarBaseColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-darkshadow-color") {=0A= e.style.scrollbarDarkShadowColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-face-color") {=0A= e.style.scrollbarFaceColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-highlight-color") {=0A= e.style.scrollbarHighlightColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-shadow-color") {=0A= e.style.scrollbarShadowColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-track-color") {=0A= e.style.scrollbarTrackColor =3D value;=0A= }=0A= else if (name =3D=3D "float") {=0A= e.style.styleFloat =3D value;=0A= }=0A= else if (name =3D=3D "table-layout") {=0A= e.style.tableLayout =3D value;=0A= }=0A= else if (name =3D=3D "text-align") {=0A= e.style.textAlign =3D value;=0A= }=0A= else if (name =3D=3D "text-align-last") {=0A= e.style.textAlignLast =3D value;=0A= }=0A= else if (name =3D=3D "text-autospace") {=0A= e.style.textAutospace =3D value;=0A= }=0A= else if (name =3D=3D "text-decoration") {=0A= e.style.textDecoration =3D value;=0A= }=0A= else if (name =3D=3D "textDecorationBlink") {=0A= e.style.textDecorationBlink =3D value;=0A= }=0A= else if (name =3D=3D "textDecorationLineThrough") {=0A= e.style.textDecorationLineThrough =3D value;=0A= }=0A= else if (name =3D=3D 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"vertical-align") {=0A= e.style.verticalAlign =3D value;=0A= }=0A= else if (name =3D=3D "visibility") {=0A= e.style.visibility =3D value;=0A= }=0A= else if (name =3D=3D "white-space") {=0A= e.style.whiteSpace =3D value;=0A= }=0A= else if (name =3D=3D "width") {=0A= e.style.width =3D value;=0A= }=0A= else if (name =3D=3D "word-break") {=0A= e.style.wordBreak =3D value;=0A= }=0A= else if (name =3D=3D "word-spacing") {=0A= e.style.wordSpacing =3D value;=0A= }=0A= else if (name =3D=3D "word-wrap") {=0A= e.style.wordWrap =3D value;=0A= }=0A= else if (name =3D=3D "writing-mode") {=0A= e.style.writingMode =3D value;=0A= }=0A= else if (name =3D=3D "z-index") {=0A= e.style.zIndex =3D value;=0A= }=0A= else if (name =3D=3D "zoom") {=0A= e.style.zoom =3D value;=0A= }=0A= }=0A= catch (e) {=0A= /* ignore error on attempt to set e.style.[property] */=0A= }=0A= }=0A= }=0A= }=0A= ------=_NextPart_000_0000_01C85F7F.F9B604C0 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://molinterv.aspetjournals.org/javascript/entrez/callback.js /************************************************************************= *****=0A= * javascript/entrez/callback.js=0A= *=0A= * Entrez Linking callback to populate content box.=0A= *=0A= * Copyright 2006 Board of Trustees of the Leland Stanford Junior = University.=0A= = *************************************************************************= ***/=0A= =0A= /*=0A= * Execute callback to fill content box with Entrez Linking information.=0A= */=0A= function entrez_callback(pmid, callback_url) {=0A= /*=0A= * MSIE 5.5 and below have issues with the JavaScript=0A= * used for Entrez Linking. For now we have to disable=0A= * the callback until we can track down a proper fix=0A= * (or everybody sanely upgrades to version 6 or 7!).=0A= */=0A= if (navigator) {=0A= var appname =3D navigator.appName;=0A= if (appname =3D=3D "Microsoft Internet Explorer") {=0A= var userAgent =3D navigator["userAgent"];=0A= var s =3D "MSIE ";=0A= var n =3D -1; =0A= if ((n =3D userAgent.indexOf(s)) !=3D -1) {=0A= var v =3D parseFloat(userAgent.substring(n+s.length));=0A= if (v < 6) {=0A= return;=0A= }=0A= }=0A= }=0A= }=0A= =0A= /*=0A= * Acquire table row element to update, initiate callback=0A= * to update table with Entrez Links.=0A= */=0A= var tr =3D document.getElementById('entrez_callback_'+pmid);=0A= if (!tr) {=0A= return;=0A= }=0A= var req =3D new XMLHttpRequest();=0A= if (!req) {=0A= return;=0A= }=0A= req.onreadystatechange =3D function() {=0A= if (req.readyState =3D=3D 4 && (req.status =3D=3D 200 || req.status = =3D=3D 304)) {=0A= var src =3D req.responseXML.documentElement;=0A= var dst =3D document.createDocumentFragment();=0A= for (var i =3D 0; i < src.childNodes.length; i++) {=0A= copy_xml_to_html(src.childNodes[i], dst);=0A= }=0A= var tbl =3D tr.parentNode;=0A= tbl.replaceChild(dst, tr);=0A= }=0A= }=0A= req.open('GET', callback_url, true);=0A= req.send(null);=0A= }=0A= ------=_NextPart_000_0000_01C85F7F.F9B604C0--