From: Subject: MicroRNA signatures in human cancers : Article : Nature Reviews Cancer Date: Thu, 8 May 2008 13:49:51 +0200 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_01AB_01C8B112.634FA1F0" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.5512 This is a multi-part message in MIME format. ------=_NextPart_000_01AB_01C8B112.634FA1F0 Content-Type: text/html; charset="iso-8859-1" Content-Transfer-Encoding: quoted-printable Content-Location: http://www.nature.com/nrc/journal/v6/n11/full/nrc1997.html MicroRNA signatures = in human cancers : Article : Nature Reviews Cancer

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Nature Reviews Cancer = 6, 857-866 = (November 2006) | = doi:10.1038/nrc1997

MicroRNA signatures in human cancers

George A. Calin1<= /A>=20 & Carlo M. Croce1<= /A> =20 Ab= out the=20 authors

T= op of page

Abstract

MicroRNA (miRNA) alterations are involved in the = initiation and=20 progression of human cancer. The causes of the widespread differential=20 expression of miRNA genes in malignant compared with normal cells can be = explained by the location of these genes in cancer-associated genomic = regions,=20 by epigenetic mechanisms and by alterations in the miRNA processing = machinery.=20 MiRNA-expression profiling of human tumours has identified signatures = associated=20 with diagnosis, staging, progression, prognosis and response to = treatment. In=20 addition, profiling has been exploited to identify miRNA genes that = might=20 represent downstream targets of activated oncogenic pathways, or that = target=20 protein-coding genes involved in cancer.

Cancer is a complex genetic disease involving structural = and=20 expression abnormalities of both coding and non-coding genes. For almost = three=20 decades, the alteration of protein-coding oncogenes and/or = tumour-suppressor=20 genes (for reviews see Refs 1=96= 3)=20 have been thought to be the causes of tumorigenesis. With the discovery = in the=20 past few years of thousands of genes that produce non-coding RNA (ncRNA) = transcripts with no significant open reading frame, it has become = evident that=20 the genomic complexity of the cancer cell is far greater than expected. = Although=20 the identification of long ncRNAs was quite a slow process owing to the=20 difficulties of experimentally cloning non-coding cDNAs of tens or = hundreds of=20 thousands of bases4<= /A>,=20 the cloning of small members of the ncRNA class was recently extremely=20 successful5,= =20 6,= =20 7,= =20 8,= =20 9<= /A>.

These microRNAs (miRNAs) are small non-coding RNAs of = 20=9622=20 nucleotides, typically excised from 60=96110 nucleotide foldback RNA = precursor=20 structures (for reviews see Refs 1= 0=9612).=20 The biogenesis of miRNAs involves a complex protein system, including = members of=20 the Argonaute family, Pol II-dependent transcription and the RNase IIIs = Drosh= a=20 and Dicer= 1= 3.=20 MiRNAs are involved in crucial biological processes, including = development,=20 differentiation, apoptosis and proliferation1= 1,=20 1= 4,=20 through imperfect pairing with target messenger RNAs (mRNAs) of = protein-coding=20 genes and the transcriptional or post-transcriptional regulation of = their=20 expression1= 5,=20 1= 6,=20 1= 7.=20 The number of human miRNAs reported so far (the July 2006 release of miRBase, at the = Sanger=20 Institute1= 8)=20 is in excess of 450, twice as many as initial calculations = indicated1= 9,=20 and more than1,000 predicted miRNA genes2= 0,=20 2= 1,=20 2= 2=20 are awaiting experimental confirmation.

Initially identified in B-cell chronic lymphocytic = leukaemia (CLL)<= A=20 href=3D"http://www.nature.com/nrc/journal/v6/n11/full/nrc1997.html#B23">2= 3,=20 changes in the expression level of miRNAs have subsequently been = detected by=20 different groups in many types of human tumours (several reviews give = various=20 details of the link between miRNAs and cancer, such as Refs 2= 4=9634).=20 MiRNAs have been proposed to contribute to oncogenesis because they can = function=20 either as tumour suppressors (as is the case for miR-15a=20 and miR-16-1) or oncogenes (as is the case for = miR -155or members of the miR-17=9692=20 cluster). The genomic abnormalities found to influence the activity of = miRNAs=20 are the same as those previously described for protein-coding genes, = such as=20 chromosomal rearrangements, genomic amplifications or deletions and = mutations.=20 In a specific tumour, abnormalities both in protein-coding genes and = miRNAs can=20 be identified2= 9.=20 Homozygous mutations or the combination of deletion plus mutation in = miRNA genes=20 are rare events3= 5,=20 and the functional consequences of heterozygous sequence variations of = miRNAs in=20 human cancers have not been identified3= 6.=20 Furthermore, the role of polymorphisms in the complementary sites of = target=20 mRNAs in cancer patients3= 7=20 or individuals with a predisposition to other hereditary diseases3= 8=20 have just started to be understood. Therefore, at the present time, the = main=20 mechanism that underlies changes in the function of miRNAs in cancer = cells seems=20 to be aberrant gene expression, characterized by abnormal levels of = expression=20 for mature and/or precursor miRNA sequences compared with the = corresponding=20 normal tissues.

MiRNA=20 profiling of tumour versus normal tissues

Oligonucleotide miRNA microarray analysis is the most = commonly=20 used high-throughput technique for the assessment of cancer-specific = expression=20 levels for hundreds of miRNAs in a large number of samples (for reviews = see Refs=20 1= 3,3= 9,4= 0).=20 Another method to determine miRNA expression levels involves the use of = a bead-based=20 flow-cytometric technique4= 1=20 (Fi= g.=20 1). Other developments include quantitative real-time PCR for = precursor=20 miRNAs4= 2=20 or active miRNAs4= 3,=20 4= 4,=20 miRAGE =97 a genome-wide miRNA analysis with serial analysis of gene = expression=20 (SAGE)4= 5=20 =97 or the high-throughput array-based Klenow enzyme (RAKE) assay4= 6.=20 Each of these techniques has its strengths and weaknesses, but after = several=20 years of study using these technologies, and the analysis of more than = 1,000=20 primary tumours, common characteristics of miRNA deregulation in human = tumours=20 have emerged.

Figure=20 1 | Principles of microarray technology and = bead-based flow cytometry used for microRNA profiling.

The process of microarray-based microRNA (miRNA) profiling, which was = initially developed by Liu et al.3= 9,=20 is shown as described in most profiling studies on primary tumours. The=20 bead-based miRNA profiling technique was developed by Lu et=20 al4= 1.=20 Both strategies involve four main steps (shown on the left side): target = labelling, DNA=96DNA hybridization, staining and signal detection. The = different=20 replicates of the spots on the glass slide represent different = oligonucleotide=20 sequences corresponding to sequences from the precursor miRNA or active = miRNA=20 molecule, whereas the different beads are marked with different = fluorescent tags=20 (each colour represents a specific miRNA using up to 100 different = colours). The=20 main advantage of the microarray-based miRNA profiling technique is the = high=20 standardization of the procedure (which enables tens of samples to be = processed=20 in parallel), whereas the main advantage of the bead-based miRNA = profiling is=20 that, because hybridization takes place in solution (instead of a glass=20 surface), it offers a higher specificity for closely related miRNAs. The = described polyacrylamide gel (PAA) fractionation is an additional = processing=20 step in the bead-based system. This figure is modified with permission = from Ref.=20 4= 9=20 =A9 (2004) National Academy of Science, USA.


MiRNA-expression profiles classify human = cancers. To=20 date, every type of tumour analysed by miRNA profiling has shown = significantly=20 different miRNA profiles (for mature and/or precursor miRNAs) compared = with=20 normal cells from the same tissue (Ta= ble=20 1, Fi= g.=20 2). Two large profiling studies using various tumours and two = distinct=20 technologies to investigate genome-wide miRNA expression have been=20 published4= 1,=20 4= 7.=20 Lu et al. developed a bead-based flow-cytometric = profiling=20 technology that is highly specific. They performed a systematic analysis = of 334=20 leukaemias and solid cancers, and found that miRNA-expression profiles = classify=20 human cancers according to the developmental lineage and differentiation = state=20 of the tumours. Furthermore, they observed patterns of gene expression = for each=20 type of tumour that reflected distinct mechanisms of transformation. For = example, the acute leukaemia samples were divided into three clusters = that=20 corresponded to the BCR=96ABL, TEL-AML1 (acute myeloid leukeamia 1, also known as = runt-related=20 transcription factor 1; RUNX1) and MLL (mixed lineage leukaemia) rearrangements, = respectively.=20 An interesting observation was that miRNA expression in tumour samples = seems=20 globally lower than in normal tissue, as 129 out of 217 measured miRNAs = showed=20 this pattern of expression. One explanation could be that global miRNA = levels=20 reflect the state of cellular differentiation. In fact, several = profiling=20 studies, including those by Lu et al.4= 1=20 and by Garzon et al.4= 8,=20 showed that distinct miRNA profiles characterize different = haematopoietic=20 differentiation stages.

Figure=20 2 | Examples of microRNA profiles in human = solid and=20 liquid cancers.

The general consensus is that profiling enabled the signatures that = are=20 associated with diagnosis =97 the process of identifying the nature of a = disease =97=20 and prognosis =97 a prediction of the probable course and outcome of a = disease =97=20 to be defined. a | The profiling of 836 solid = tissue and=20 haematological primary cancers (and corresponding normal tissues = samples) using=20 the microarray technology (as described in Ref. 3= 9)=20 and reported to date is shown. The exact number of analysed samples from = each=20 tumour type (corresponding normal controls included) is shown. The data = were=20 collected from the references cited in the text. b | The=20 common expression signature in six solid cancers indicates common = altered=20 regulatory pathways involving the same microRNA (miRNA) genes (named on = the=20 right side of the dendrogram)4= 7.=20 These miRNAs could be considered as 'cancer markers', because variations = of=20 their expression could identify the cancerous state. c |=20 The distinct clusters identified by miRNA profiles in chronic = lymphocytic=20 leukaemia (CLL)4= 9=20 were further confirmed to be associated with prognostic factors and = disease=20 progression3= 5.=20 Note also that various normal haematopoietic samples cluster in a group = that is=20 distinct from all CLL samples. MNC, mononuclear cells; Ly, B = lymphocytes; CD5, a=20 subset of B lymphocytes largely accepted to represent the equivalent of=20 malignant cells in CLL9= 3.=20 Panel b is modified with permission from Ref. 4= 7 =A9=20 (2006) National Academy of Science, USA.


Table=20 1 | Facts about microRNA-expression = profiling in=20 human cancers

Using a large-scale microarray analysis of 540 samples, = including=20 363 from six of the most frequent human solid tumour types and 177 = normal=20 controls, Volinia et al. also found that cancer = cells=20 showed distinct miRNA profiles compared with normal cells4= 7=20 (Fi= g.=20 2). Out of the 228 miRNA genes analysed, 36 were overexpressed and = 21 were=20 downregulated in cancer cells versus normal cells. Hierarchical=20 clustering analyses showed that this miRNA signature enabled the = tumour=20 samples to be grouped on the basis of their tissue of origin. Several = other=20 genome-wide profiling studies have been performed on various cancer = types,=20 including CLL4= 9,=20 breast=20 cancer5= 0,=20 glioblastoma5= 1,=20 thyroid = papillary=20 carcinoma3= 7,=20 hepatocellular=20 carcinoma5= 2,=20 lung = cancer5= 3,=20 colon = cancer4= 5=20 and endocrine=20 pancreatic tumours5= 4.=20 For example, in a study of 104 matched pairs of primary cancerous and=20 non-cancerous lung tissue, Yanaihara et al. found = a set of=20 43 differentially expressed miRNAs; 28 were downregulated and 15 were=20 overexpressed in the tumour cells5= 3.=20 Murakami et al., using a microarray platform, = found that in=20 hepatocellular carcinoma five out of eight differentially expressed = miRNAs were=20 downregulated in cancer cells5= 2.=20 All these studies support the same view: the alterations seen in cancer = cells=20 that express miRNAs consist of both overexpressed and downregulated=20 microRNAs.

One key issue is the level of differential expression = that can be=20 considered biologically significant. 10=9620-fold differences in miRNA = expression=20 levels have been found to be important in papillary thyroid = carcinomas3= 7,=20 however, we share the view that much smaller changes (less than twofold) = could=20 also be significant3= 5,=20 4= 9,=20 5= 0,=20 5= 1.=20 This reflects the fact that specific miRNAs target many genes and, as a = large=20 number of miRNAs seem to be altered in human tumours, one can postulate = that the=20 disruption of two or more cooperating oncogenes and/or tumour = suppressors is a=20 likely event2= 9.=20 An opposite view is that in some cases the described changes in miRNA = expression=20 between normal and tumour cells could be biologically irrelevant: the = main=20 interactions with various targets could have antagonistic rather than=20 accumulating biological consequences (for example, the repression of = both=20 pro-apoptotic and anti-apoptotic genes).

Common miRNA genes are differentially expressed in = various=20 cancers, suggesting common altered regulatory pathways. = Assembling the=20 statistical analyses of microarray data obtained by two different = methods, significance=20 analysis of microarrays (SAM) and prediction=20 analysis of microarrays (PAM) from six solid tumours (lung, breast, = colon,=20 gastric and prostate carcinomas and endocrine pancreatic tumours), = Volinia et al. described a common signature composed of 21 = miRNAs=20 differentially expressed in at least three tumour types4= 7.=20 At the top of the list were miR-21, which was = overexpressed=20 in six types of cancer cells, and miR-17-5p and = miR-191, which were overexpressed in five. As the = embryological=20 origin of the analysed tumours was different, the significance of such = findings=20 could be that these common miRNAs participate in fundamental signalling = pathways=20 altered in many types of tumour. Supporting the function of these genes = in=20 tumorigenesis, it was found that the predicted targets for the = differentially=20 expressed miRNAs are significantly enriched for those that target known = tumour=20 suppressors and oncogenes4= 7.=20 Furthermore, miR-21, the only miRNA overexpressed = in all=20 six types of cancer analysed, and which is also overexpressed in=20 glioblastomas5= 5,=20 5= 1=20 and cholangiocarcinomas5= 6,=20 was shown by Meng et al. to directly target the = tumour=20 suppressor PTEN in cholangiocarcinoma cells. PTEN=20 encodes a phosphatase that can inhibit growth and/or survival pathways, = and its=20 function is altered in advanced tumours of various types, including = breast,=20 lung, gastric and prostate5= 7.=20 In all these tumours, the frequency of loss-of-heterozygosity (LOH) at = the PTEN locus is relatively small (less than 10%), leading = to the=20 speculation that if PTEN is shown to be a target = of miR-21 in the other tumour types then this could be the = main=20 mechanism of PTEN inactivation. Furthermore, = knockdown of=20 miR-21 in cultured glioblastoma cells triggers = apoptosis by=20 a caspase-dependent mechanism5= 5.=20 All these data suggest that miR-21 is an = anti-apoptotic and=20 pro-survival factor.

Further examples of miRNAs that are frequently = deregulated in=20 cancer and target tumour suppressors and/or oncogenes come from an = elegant study=20 performed in the laboratory of F. Slack, which clearly showed that Ras = oncogenes=20 are regulated by the let-7 miRNA family5= 8.=20 Another example of fine functional dissection of a miRNA pathway comes = from the=20 study of Voorhoeve et al., which shows the = oncogenic=20 function of miR-372 and miR-373 in=20 testicular germ-cell tumours of adolescents and adults. These miRNAs = facilitate=20 the proliferation and transformation of cells that harbour both = oncogenic HRAS and functional wild-type TP53 (Ref. 5= 9).

Additional support for the function of miRNAs in human = cancers has=20 come from transgenic and knockout mouse models, which have been = important for=20 understanding the tumorigenic pathways in which deregulated miRNAs are = involved.=20 He et al. showed that enforced expression of the = miR-17-92 cluster accelerated MYC-induced=20 lymphomagenesis6= 0,=20 suggesting the oncogenic potential of some of the miRNAs that are = overexpressed=20 in B-cell lymphoma samples owing to genomic amplification. Recently, = Costinean=20 et al. produced the first transgenic mouse that=20 specifically overexpresses a miRNA gene, miR-155, = in B=20 cells6= 1.=20 This gene is overexpressed in various types of B-cell malignancy6= 2,=20 6= 3,=20 6= 4.=20 The transgenic mice developed polyclonal pre-leukaemic pre-B-cell = proliferation=20 followed by B-cell malignancy. These findings suggest that miR-155 deregulation could be an early event in = oncogenesis that=20 needs additional genetic alterations for the development of the fully = malignant=20 phenotype6= 1.=20 The same was also shown for the MYC oncogene, = which is=20 activated by chromosomal translocation in human B-cell neoplasia6= 5.=20 In an avian model, oncogenic cooperation in lymphomagenesis and=20 erythroleukaemogenesis between the non-protein-coding BIC gene, which was later shown to include the = miR-155 sequence, and MYC has been observed6= 6,=20 indicating that MYC and BIC might cooperate in = human=20 tumours. A microarray study of the malignant B cells isolated from miR-155 transgenic mice compared with non-transgenic = controls=20 resulted in a complete list of the deregulated protein-coding genes in = these=20 cells, and therefore potentially identified direct and indirect targets = of miR-155. Of particular note, the expression of 16 = different=20 miRNAs was altered in pre-malignant cells from the transgenic = mice6= 1,=20 supporting the view that complex relationships exist between = protein-coding=20 genes and miRNAs that affect specific signalling pathways and, probably, = direct=20 or indirect interactions exist between miR-155 = and some of=20 the other deregulated miRNAs.

Onco-miRNAs and tumour-suppressor-miRNAs: two = different=20 views of the same gene? Genome-wide expression studies also = offer=20 intriguing new perspectives on the involvement of miRNAs in human = cancer. The=20 first perspective comes from the studies performed by O'Donnell et=20 al. on cell models, which showed that in the human B-cell line = P493-6,=20 which overexpresses MYC, miRNA members of the = miR-17=9692 locus have tumour-suppressor activity = because their=20 expression decreases the expression of E2F1, and so inhibits = MYC-mediated=20 cellular proliferation6= 7.=20 However, a different perspective is offered by the results from B-cell=20 lymphomas. The same cluster of miRNAs function as a potential oncogene = by=20 cooperating with MYC and blocking = apoptosis6= 0.=20 One possible explanation is that the same miRNA can participate in = distinct=20 pathways, having different effects on cell survival, growth and = proliferation=20 that are dependent on the cell type and the pattern of gene expression. = The=20 combinatorial nature of miRNA=96mRNA interactions means that the same = miRNA could=20 have different targets and the same mRNA could be targeted by different = miRNAs=20 in different cell types. For example, Cimmino et = al. found=20 that in leukaemic cells with a deletion of the locus on chromosome 13 = that=20 contains miR-15a and miR-16-1 miRNA=20 genes, apoptosis is induced as a result of the exogenous re-expression = of these=20 miRNAs. MiR-15a and miR-16-1 interact=20 with the anti-apoptotic gene BCL2 when added back to these cells, however, a = similar=20 response is not seen in 293 fetal kidney cells, which normally express = miR-15a, miR-16-1 and BCL2 (Ref. = 6= 8).=20 Therefore, it is probable that the loss of miR-15a and=20 miR-16-1 induces the overexpression of BCL2 in B = cells,=20 giving rise to B-cell malignancies. In a different tissue, the = overexpression of=20 miR-15a and miR-16-1 might = cause the=20 loss of expression of a tumour suppressor that is important for growth = control=20 and/or apoptosis. In support of this hypothesis are reports that show = that,=20 althoughmiR-15a and miR-16-1 are=20 downregulated in B lymphocytes from patients with CLL2= 3,=20 the same miRNAs are overexpressed in endocrine pancreatic tumours of=20 neuroectodermal origin4= 7,=20 5= 4.

The second perspective comes from the finding that the = consistent=20 abnormal expression of the precursor miRNAs, but not the active = molecules, can=20 be identified in various types of cancer4= 2,=20 4= 7,=20 4= 9,=20 5= 2,=20 5= 3.=20 Yanaihara et al. found that in their 43 miRNA = gene=20 signature, seven precursor miRNAs were differentially expressed in = matched lung=20 cancer versus normal lung tissue. In addition, Jiang et=20 al., using a real-time PCR method specifically designed for = precursor=20 profiling4= 2,=20 found significant variations in the expression of nine precursors in a = panel of=20 32 cancer cell lines6= 9.=20 These data raise the possibility that the 'non-active' part of the miRNA = molecule could have 'independent' and as yet unknown functions that = could be=20 important in tumorigenesis. Putative interaction partners might be the=20 RNA-binding proteins (such as the heterogenous nuclear = ribonucleoproteins,=20 hnRNPs), a class of proteins that are abnormally expressed in all types = of human=20 cancers7= 0.=20 These proteins bind RNA molecules to regulate their stability, and = therefore it=20 is possible that in the sequence of several precursor miRNAs such=20 hnRNP-interacting motifs exist. However, there are no data so far that = directly=20 attribute regulatory function to a miRNA precursor.

Causes of = abnormal miRNA=20 expression

The causes of the widespread disruption of miRNA = expression in=20 tumour cells are only partially known and, very probably, various = abnormalities=20 in each tumour could contribute to the global miRNA-expression profile. = So far,=20 at least three different mechanisms (that could function independently = or=20 together) have been described. Such alterations correlate with and could = explain=20 the abnormal expression of specific miRNA genes.

The location of miRNAs at cancer-associated = genomic regions=20 (CAGRs). Soon after the identification of hundreds of new = members of the=20 miRNA family, it was shown that more than half of the known human miRNAs = reside=20 in particular genomic regions that are prone to alteration in cancer=20 cells7= 1.=20 Such regions include minimal regions of LOH, which are thought to = harbour=20 tumour-suppressor genes, minimal regions of amplification, which might = contain=20 oncogenes, common breakpoint regions in or near possible oncogenes or=20 tumour-suppressor genes and fragile sites (FRA). FRA are preferential = sites of=20 sister-chromatid exchange, translocation, deletion, amplification or = integration=20 of plasmid DNA, and tumour-associated viruses such as human papilloma = virus=20 (HPV). Of note, miRNA loci were significantly associated with insertion = sites of=20 tumorigenic HPV strains linked to endometrial cancers, leading to the=20 possibility that viral insertions in the human genome might disturb = small=20 ncRNAs7= 1.

Several reports show that this computationally = identified=20 association is significant for miRNA expression. Zhang et=20 al. showed, by using high-resolution array comparative genomic=20 hybridization (aCGH), that the miRNA loci have a high frequency of = genomic=20 alteration in human cancers7= 2.=20 Furthermore, about 75% of the miRNAs analysed showed concordance between = mature=20 miRNA levels and DNA copy number, and, importantly, their genomic data = were in=20 agreement (81% concordance for amplified and/or overexpressed miRNAs, = and 60%=20 concordance for deleted and/or downregulated miRNAs, respectively) with = the=20 expression data published with an independent set of breast cancers by = Iorio=20 et al5= 0.=20 Other examples of concordance by two independent reports, involve two = miRNA=20 clusters located at chromosome 13 (Fi= g.=20 3). The cluster miR-15a=96miR-16-1, located = at 13q14.3 in=20 a region deleted in B-cell chronic lymphocytic leukaemia and pituitary = adenomas=20 was shown by northern blots to be downregulated in most patients with = these=20 tumours2= 3,=20 7= 3.=20 The cluster miR-17=9692, located at chromosome = 13q31, in a=20 region amplified in B-cell lymphomas and lung cancers, was found to be=20 overexpressed using microarray studies by He et = al.5= 9,=20 and by Hayashita and colleagues and Tagawa and Seto using northern blot = analyses=20 in lung cancers and malignant lymphomas, respectively7= 4,=20 7= 5.

Figure=20 3 | Chromosomal alterations at microRNA=20 loci.

The main chromosomal alterations at microRNA (miRNA) loci, loss of=20 heterozygosity and amplification, are identified at two separate regions = of=20 chromosome 13. a | Shows the 13q14.3 deletion = found in=20 chronic lymphocytic leukaemia (CLL) as described in Ref. 2= 3.=20 The downregulation of miR-15a and miR-16-1 induces overexpression of the anti-apoptotic = BCL2=20 protein in leukaemia cells6= 8.=20 The genomic regions are not drawn to scale, and the cluster on 13q13 is=20 simplified. The functional consequences are shown on the right of the = figure.=20 b | Shows the 13q13 amplification identified in = high-grade=20 lymphomas, as described in Ref. 7= 4.=20 The functional consequences are presented on the right side of the = figure.The=20 overexpression of miR-17-5p and miR-20a from the same cluster induces the = downregulation of the=20 transcription factor E2F1. It was also shown that the MYC=20 oncogene increases the expression of the miR-17=9692 cluster=20 and activates E2F1 (Ref. 6= 7),=20 and that the MYC-activated cluster increases tumour angiogenesis= 113.=20 Therefore, the overexpression of the miR-17=9692 = cluster is=20 correlated with hyperproliferation, the blocking of apoptosis and=20 neovascularisation. The arrows and the bars represent stimulatory and = inhibitory=20 signals, respectively. Orf, open reading frame.


Epigenetic regulation of miRNA expression. = DNA=20 hypomethylation, CpG island hypermethylation and histone-modification = losses=20 represent epigenetic markers of malignant transformation7= 6.=20 Therefore, several groups have investigated whether such events also = affect=20 miRNA expression. Scott et al. showed that in = breast cancer=20 cells histone deacetylase inhibition is followed by the extensive and = rapid=20 alteration of miRNA levels7= 7.=20 Furthermore, Saito et al. found that the combined = treatment=20 of human bladder cancer cells with 5-aza-2'-deoxycytidine (5-Aza-CdR) = and the=20 histone deacetylase (HDAC) inhibitor 4-phenylbutyric acid (PBA) has a=20 significant effect on the expression of miRNAs7= 8.=20 Seventeen miRNAs (out of 313 screened by a microarray assay) were = upregulated=20 more than threefold, and miR-127 was the most=20 differentially expressed. This miRNA is located in a CpG island at = chromosome=20 14q32, a region that is involved in several types of translocations = identified=20 in haematological cancers and deleted by LOH in solid tumours7= 1.=20 Furthermore, the combined treatment was accompanied by a decrease in DNA = methylation and an increase in active histone markers around the = transcription=20 start site of miR-127. The investigators showed = that miR-127 can translationally repress the transcription = of the=20 zinc-finger repressor BCL6 (Ref. 7= 8).=20 Therefore, the induction of miR-127 by 5-Aza-CdR = and PBA=20 treatment in cancer cells might have an anticancer effect by = downregulating=20 anti-apoptotic factors, such as BCL6, and = inducing=20 apoptosis7= 8.

The effects of DNA demethylation and histone = deacetylation on=20 miRNA expression could be tissue-specific, as the treatment of non-small = cell=20 lung cancer (NSCLC) cells with demethylating agents and/or HDAC = inhibitors does=20 not have a significant influence on miRNA expression3= 6,=20 5= 3.=20 Supporting this assumption is the fact that miR-127 was not=20 found to be expressed in normal lung cells by microarray analysis3= 9,=20 and was not differentially expressed between cancerous and non-cancerous = lung=20 tissue5= 3.

Abnormalities in miRNA-processing genes and=20 proteins. The protein machinery that is involved in the = biogenesis of=20 miRNAs is complex. Theoretically, alterations of these proteins (for a=20 comprehensive description see Ref. 3= 4)=20 should have dramatic effects on miRNA expression. In fact, it was shown = that a=20 failure of the Drosha processing step could explain the downregulation = of miRNAs=20 observed in primary tumours7= 9.=20 Furthermore, in a large study of patients with NSCLC, Karube et=20 al. found that the expression levels of Dicer, but not Drosha, = were=20 reduced in a fraction of lung cancers. This correlated with reduced=20 post-operative survival and poor tumour differentiation status, both of = which=20 are associated with a poor prognosis8= 0,=20 8= 1.=20 A similar correlation with reduced survival in patients with NSCLC was = found for=20 the expression of miR-155 and let-7=20 (Refs 5= 3,8= 2).=20 These results suggest that patient survival could be a consequence of = both=20 aberrant miRNA processing and aberrant expression. The conditional = knockout=20 Dicer mouse has developmental defects, including lung epithelium=20 abnormalities8= 1,=20 8= 3,=20 but no reports on tumour formation are available, and Dicer-deficient = embryonic=20 stem cells injected into nude mice do not form tumours8= 4.

The influence of genetic abnormalities in = small-RNA-processing=20 proteins could have a more significant influence than is recognized at = present.=20 Recently, a new class of germline-specific small ncRNAs (26=9631 = nucleotides=20 long), named PIWI-interacting RNAs (piRNAs), was discovered and found to = bind=20 the testis-specific murine PIWI protein orthologues8= 5,=20 8= 6,=20 8= 7,=20 8= 8,=20 8= 9.=20 Intriguingly, the human homologue of this subgroup of the Argonaute = proteins,=20 named HIWI,=20 is located in the genomic region 12q24.33 that is genetically linked = with the=20 development of testicular germ-cell tumours in adolescents and adults. = In=20 addition, HIWI is overexpressed in most = testicular=20 seminomas analysed so far9= 0.=20 The expression of HIWI in human gastric cancer is = associated with proliferating cells9= 1.=20 These correlations raise the possibility that piRNAs could represent = another=20 example of cancer-altered small ncRNAs, specifically in relation to PIWI = overexpression.

MiRNA = profiling as a=20 new clinical tool

As active players in important human oncogenic = signalling=20 pathways, miRNAs should affect cancer diagnosis and prognosis (Ta= ble=20 1). As discussed above, several reports have already shown this for = several=20 tumour types4= 9,=20 5= 0,=20 6= 2.=20 But other reports indicate that miRNA profiles can be used to diagnose = tumour=20 types that cannot be determined on the basis of tumour biopsy = samples.

MiRNA profiling as a diagnostic tool. = Metastatic=20 cancer of unknown primary site (CUP) is one of the 10 most frequent = cancer=20 diagnoses worldwide, and constitutes 3=965% of all human = malignancies9= 2.=20 Patients with CUP present with metastases (late-stage disease) without = an=20 established primary tumour (that is, a site at which the tumour has = initially=20 developed and from which it has metastasized). The study of Lu et=20 al. produces an important advance in the diagnosis of this = peculiar type=20 of cancer. Analysing 17 poorly differentiated tumours with = non-diagnostic=20 histological appearance, they showed that the miRNA-based classifier was = much=20 better at establishing the correct diagnosis of the samples than the = mRNA=20 classifier4= 1.=20 This result is exciting =97 profiling a few hundred miRNAs has a much = better=20 predictive power for CUP diagnosis than profiling several tens of = thousands of=20 mRNAs. As miRNA expression changes with differentiation, the poorly=20 differentiated tumours have lower global expression levels of miRNAs = compared=20 with well-differentiated tumours from control groups4= 1.=20 The reduced expression levels of miRNAs in poorly differentiated tumours = is=20 probably why miRNA profiling is effective in the diagnosis of CUP.

CLL is the most common adult leukaemia in the Western = world9= 3,=20 9= 4.=20 During its natural history, this indolent disease can evolve into an = aggressive=20 type of lymphoma and/or leukaemia, including Richter=20 syndrome9= 5.=20 Furthermore, as the prognosis of most cases of CLL is relatively good, = treatment=20 after diagnosis is started only if poor prognostic markers are = evident9= 6.=20 Few such markers have been found, but those that have include high = expression=20 levels of the 70 kDa zeta-associated protein (ZAP70<= /A>)=20 and an absence of mutations in the immunoglobulin heavy-chain = variable-region=20 gene (IgVH). By performing a = miRNA-profiling=20 screen on 144 patients with CLL, a unique signature of 13 miRNAs (out of = the 190=20 analysed) was shown to differentiate cases on the basis of a good or bad = prognosis, or on the presence or absence of disease progression3= 5.=20 Among them, miR-16-1 and miR-15a were=20 expressed at lower levels in patients with a good prognosis, in = agreement with=20 early reports that 13q14.3 genomic deletions at the locus that harbours = these=20 genes correlate with a favourable course of the disease9= 7.=20 Among the genes found to be downregulated in the group of patients with = a bad=20 prognosis were members of the miR-29 family. This = could be=20 explained by the recent findings that TCL1, an oncogene that is overexpressed in CLL = cells from=20 patients with aggressive disease9= 8,=20 is a target of miR-29 genes (Y. Pekarsky, = unpublished=20 observations). Therefore, these findings support a model in which two = different=20 molecular types of CLL with opposite clinical behaviours exist: one with = a good=20 prognosis and slow disease progression characterized by deletions of = chromosome=20 13q14.3 and low levels of miR-15a and miR-16-1, and a second that is aggressive and prone to = develop=20 Richter syndrome, characterized by low levels of expression of miR-29 genes and high expression levels of TCL1 = protein. As BCL2 is consistently overexpressed in CLL = samples9= 9,=20 miR-15a and miR-16-1 might = have=20 additional targets important for progression; furthermore, other = mechanisms of=20 BCL2 deregulation (such as the minority of cases that have a BCL2=20 translocation= 100),=20 and other interacting miRNAs might be involved.

MiRNA profiling as a prognostic tool. Lung = cancer is=20 the leading cause of death from cancer in males worldwide= 101.=20 Therefore, the identification of new prognostic markers (markers that = correlate=20 with disease evolution) could be a significant advance for the = identification of=20 patients that would benefit from more aggressive therapy. In univariate=20 analyses, the expression of both miR-155 = (high levels)=20 and let-7a-2 (low levels)has been shown to = correlate with=20 poor survival in 104 United States patients with lung cancer; by = multivariate=20 analyses the expression of miR-155 also = correlated with a=20 poor prognosis when all clinical variables were considered = together5= 3.=20 In an independent study of 143 Japanese patients with lung cancer, = reduced let-7 expression was found to significantly correlate = with a=20 shorter survival time after potentially curative surgery8= 2.=20 In both studies, about two thirds of the cases were adenocarcinomas (a = type of=20 NSCLC). By multivariate analyses, a low expression level of let-7 was an independent prognostic factor for the = stage of the=20 disease, and correlated with significantly shorter survival8= 2.=20 In a meta-analysis of 29 studies that investigated the role of Ras = oncogenes in=20 the survival of patients with NSCLC cancers, Ras overexpression was = associated=20 with a reduced survival time= 102.=20 As let-7 negatively regulates Ras5= 8,=20 these results show the existence of a link between let-7,=20 Ras expression and the life expectancy of patients with lung cancer.

A miRNA = perspective in=20 cancer therapy

The potential for using miRNAs in cancer is now being = explored.=20 The theoretical rationale is based on the fact that miRNAs are natural = antisense=20 interactors that regulate many genes involved in eukaryotic survival and = proliferation. Also, miRNA-expression profiles have been shown to change = during=20 treatment with gemcitabine, and the modulation of some miRNAs (including = the=20 overexpression of miR-21) increases the = sensitivity of=20 cholangiocarcinoma tumour cells to this chemotherapeutic agent in=20 vitro5= 6.=20 Such results offer the experimental bases for the use of miRNAs as = therapeutic=20 targets. The development of modified miRNA molecules with longer in vivo half lives and efficiency, such as the locked = nucleic=20 acid (LNA)-modified oligonucleotides= 103,=20 the anti-miRNA oligonucleotides (AMOs)= 104=20 and the 'antagomirs'= 105=20 is the first step towards bringing these fundamental research advances = into=20 medical practice. Future in vivo experiments of = miRNA=20 transgenics and knockouts will offer valuable information about safety = and=20 efficacy.

A new form of = cancer=20 predisposition?

The expression levels of many genes show abundant = natural=20 variation in species that range from yeasts to humans. This trait, = termed 'gene=20 expression phenotype' has an inherited component, which was shown to be = present=20 through familial aggregation studies in humans and simple segregation = studies in=20 yeast= 106,=20 = 107.=20 Cheung et al., using microarray studies of human=20 lymphoblastoid cells, identified a set of genes with a high degree of = variation=20 in expression levels between individuals= 106.=20 The variation in expression among unrelated individuals was 3=9611 times = greater=20 than between monozygotic twins, and the variance among siblings was = 2=965 times=20 greater than that between twins. These data suggest that genetic = differences=20 among individuals contribute to the variation in expression phenotype = for some=20 genes= 106.=20 Because of the particular way in which miRNAs function =97 by targeting = a number=20 of functionally important protein-coding genes =97 we propose that = variations in=20 the expression and/or sequence of miRNAs, if inherited in the germline, = could be=20 important in cancer predisposition by affecting the levels of expression = of=20 tumour-suppressor genes or oncogenes (Bo= x1,=20 Fi= g.=20 4).

Figure=20 4 | The involvement of microRNAs in cancer=20 predisposition.

MicroRNA (miRNA) alterations could predispose to cancer development. = Here we=20 propose a model in which structural and/or expression abnormalities of = miRNAs in=20 the germline could represent an inherited predisposing event (blue = person). For=20 simplicity, we present only the situation of gene deletion, but all = other types=20 of loss-of-function or gain-of-function mutations described for = protein-coding=20 genes can be involved in the disruption of miRNA expression. To develop = a=20 malignant tumour, a second genetic event has to occur in a somatic cell = (green=20 people): this can be an alteration in a protein-coding gene, or a 'hit' = in=20 another miRNA (shown here as amplification). The consequences are on the = levels=20 of expression of various target mRNAs: the overexpression of target = oncogenes=20 (red protein) in the case of miRNA deletion, and the downregulation of = target=20 tumour-suppressor genes in the case of miRNA amplification (purple = protein).


Bo= x 1 |=20 Putative mechanisms of cancer predisposition by microRNA = alterations
  • Fu= ll box=20

A significant proportion of patients with cancer are = members of=20 kindreds that develop the same type of cancer without a specific = Mendelian=20 inheritance, or that develop different types of cancers. This type of = familial=20 predisposition represents an important medical problem, and the = identification=20 of the predisposing gene(s) could offer early diagnosis and potential = cancer=20 prevention. Despite various efforts, the basis of this type of = predisposition is=20 largely unknown. A clear example comes from cases of familial CLL: = although rare=20 mutations in the ataxia telangiectasia mutated (ATM)= 108,=20 = 109=20 or ARLTS1 genes= 110,=20 as well as in the miR-15a and miR-16-1=20 transcripts3= 5,=20 have been identified in families with two or more members with CLL, the = culprit=20 genes are still unknown for most cases.

In the United States, prostate cancer affects = African-American=20 males at a rate that is disproportionate with the number of White males=20 diagnosed with the disease, and genetics could have an important role, = but no=20 target genes have been identified after decades of research= 111.=20 We analysed miRNA expression levels in a pilot set of 18 samples of = normal=20 prostate tissue derived from two distinct human populations, 10 = Caucasians from=20 the United States and 8 African-Americans collected by the US National = Cancer=20 Institute Cooperative Prostate Cancer Tissue Resources. Using two = different=20 methods of microarray analyses, PAM and SAM, we identified both = significantly=20 overexpressed and downregulated miRNAs in the African-American = individuals=20 compared with the Caucasian individuals (G.A.C., C.M.C., S. Volinia and = S. Ambs,=20 unpublished data). The list of miRNAs that are at least three times=20 differentially expressed includes miR-301, miR-219, miR-26a, miR-1b-1 and miR-30c-1. = Therefore, different=20 human populations show variations in the global expression pattern of = miRNAs=20 expressed in a specific tissue. Whether such variations represent an = inherited=20 family trait remains an open question that should be solved by future = large=20 studies aimed at assessing the global miRNA expression profiles in = cancers with=20 high heritability, such as familial breast cancers or familial CLLs.

In conclusion

The question of whether miRNAs represent the 'dark side' = of cancer=20 predisposition remains to be answered, but there is no doubt that these=20 'strangers' in the genomic galaxy are involved in the regulation of = pathways=20 that are involved in the development and progression of many types of=20 tumour.

T= op of page

Acknowledgements

We want to thank the collaborators and young = investigators who=20 contributed to the microRNA work done in our laboratories. C.M.C. is = supported=20 by Program Project grants from the US National Cancer Institute, and = G.A.C. is=20 supported by a Kimmel Foundation Scholar award and by the CLL Global = Research=20 Foundation. We thank M. Negrini and M. Fabbri for their critical reading = of the=20 manuscript, and thank A. Cimmino for assistance with the figures. We = apologize=20 to many colleagues whose work was not cited owing to space = limitations.

Competing interests statement

The authors declare no competing financial=20 interests.

T= op of page

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Author affiliations

  1. Department of Molecular Virology, Immunology and Medical = Genetics=20 and Comprehensive Cancer Center, Ohio State University, Columbus, Ohio = 43210,=20 USA.

Correspondence to: Carlo M. Croce1<= /A>=20 Email: Carlo.Croce@osumc.edu

=
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