ࡱ> h j S T U V W X Y Z [ \ ] ^ _ ` a b c d e f g  Jbjbj<< ((^^ݯ/  8\{L;( !"""<#<#<#:::::::8=?:]<#8#<#<#@#:2""3R;222D# ""O62<#:22s55f /56<h;0;5,@2@52585<#<#<# :   A chapter for a book Genome-scale Pattern Analysis in the Post-ENCODE Era. An Intricate Mosaic of Genomic Patterns at Mid-range Scale Alexei Fedorov and Larisa Fedorova Department of Medicine, University of Toledo, Health Science Campus. Introduction Genomic patterns on short-range scales represent various words composed from nucleotide letters. Each of these words occurs many times within DNA sequences. The longest words, also known as pyknons, are up to 17 nucleotide-long sequences which are over-abundant in the exons and introns of humans and other mammals  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Rigoutsos et al. 2006; Tsirigos and Rigoutsos 2008). The vast majority of sequences only a little bit longer than pyknons are unique even for the large genomes of animals and plants. For example, the complete theoretical set of 20 nucleotide-long sequences is comprised of 420 different words of length 20, which is just over one trillion. More than 99% of these 20-mer oligonucleotides never occur in the entire human genome (~ 3*109 bp). Therefore, biologists frequently use 20-mer oligonucleotides as PCR primers or hybridization probes for experimental characterization of particular genomic segments. The genomic arrangement of short sequences (< 20 bp) is covered in other chapters of this book. Here we consider genomic patterns longer than 30 and up to several thousands of nucleotides to be called mid-range scale. At this mid-range, most of the sequences are unique, i.e. occur only once in the entire genome, hence, it is more appropriate to characterize or group them not by their exact sequence of nucleotides but rather by their overall nucleotide composition, such as G+C-richness, purine-richness, etc. We also distinguish mid-range genomic scales from the long-range one represented by genomic isochores reviewed elsewhere  ADDIN EN.CITE Bernardi200748048048017Bernardi, G.Molecular Evolution Laboratory, Stazione Zoologica Anton Dohrn, Villa Comunale, 80121 Naples, Italy. bernardi@szn.itThe neoselectionist theory of genome evolutionProc Natl Acad Sci U S AProc Natl Acad Sci U S A8385-90104202007/05/15AnimalsBase Composition/geneticsChromosomes/genetics*EvolutionGenetic CodeGenome/*geneticsHumansIsochores/genetics*Models, GeneticSyntenyTime FactorsVertebrates/*genetics2007May 150027-8424 (Print)17494746http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1749474618663110701652104 [pii] 10.1073/pnas.0701652104eng(Bernardi 2007). Traditionally, G+C-rich and G+C-poor isochors are considered to be from 100 kb and longer. Recently, scientists have started to describe ultra-short isochores in the range of tens of thousands of nucleotides. In order not to interfere with isochors, we limit the length of mid-range patterns by ten thousand bases. The main focus of this paper is to show that at mid-range scales, genomes of complex eukaryotes consist of a number of different patterns and are associated with unusual DNA conformations. Some of these patterns are scarcely investigated and still wait for thorough exploration and recognition. Results and Discussion DNA repeats important elements at genomic mid-range scale All eukaryotic genomes contain several extra-large words recurring many times -- so called DNA repetitive elements, the size of which are generally within mid-range scale. DNA repeats are classified into three major classes based on the molecular mechanisms of their origin and propagation: transposons, retrotransposons, and tandemly organized repeats. There is a large variety of transposons and retrotransposons that can be specific for narrow taxons (like the Alu-elements within primates), or that have a much broader representation (like the L1-repeats found in all vertebrates). We will not consider these DNA repetitive elements, but only refer a reader to several excellent, detailed reviews on their genomic organization and evolution  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Jurka et al. 2007; Eickbush and Jamburuthugoda 2008; Richard, Kerrest, and Dujon 2008). Here we concentrate only on the simple tandem repeats that exist in almost every eukaryotic organism. Our examples well illustrate the common trend for mid-range scale sequence patterns to associate with DNA conformation abnormalities or alternative 3-D structures. We begin the examination of a simple tandem repeat from a well-characterized type composed of a reiterating pentamer sequence AATGG. Our computer analysis of completely sequenced mammalian genomes demonstrates that there were 162 different loci inside the euchromatic genomic regions of humans, 24 in mouse, 14 in rat, 21 in cow, and 58 in dog that contain (AATGG)N perfect repeats, where Ne"4. These sequences are proportionally distributed between intergenic regions and introns and, often, there are up to several dozens of tandem AATGG pentamers in one locus. The location of these repeats inside genes is not evolutionarily conserved, since we have not detected their presence in the same intron of the named species. In addition, the same tandemly repeated pentamer AATGG is one of the most evolutionarily-conserved parts of a centromere, where it exists in thousands of copies and serves as an attachment point for the two sister chromosomes during mitosis  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Grady et al. 1992; Lee et al. 1997). Interestingly, under physiological conditions, this DNA-repeat comprised of at least four pentamers could exist not only as B-form Watson-Crick duplex but also in an unusual form with highly asymmetrical conformations of AATGG-strand and its complementary CCATT-strand  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Jaishree and Wang 1994; Catasti et al. 1999). Its transition from Watson-Crick duplex to single-stranded structures is facilitated by acidic pH conditions. Figure 1 demonstrates the NMR solution structure of the anti-parallel stranded non-B-form DNA duplex 5-TGGAATGGAA:TGGAATGGAA-3 created by two repetitive pentamers published by  ADDIN EN.CITE Chou199429292917Chou, S. H.Zhu, L.Reid, B. R.Howard Hughes Medical Institute, University of Washington, Seattle 98195.The unusual structure of the human centromere (GGA)2 motif. Unpaired guanosine residues stacked between sheared G.A pairsJ Mol BiolJ Mol Biol259-6824431994/12/02Adenosine/chemistryBase CompositionBase SequenceCentromere/*chemistry/geneticsComputer GraphicsDNA/*chemistryGuanosine/chemistryHumansMagnetic Resonance SpectroscopyMolecular Sequence Data*Nucleic Acid Conformation*Repetitive Sequences, Nucleic Acid1994Dec 20022-2836 (Print)7966337http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=7966337S0022-2836(84)71727-X [pii] 10.1006/jmbi.1994.1727eng(Chou, Zhu, and Reid 1994). This particular structure is also known as interdigitated or zipper-like stacking  ADDIN EN.CITE Chou200339393917Chou, S. H.Chin, K. H.Wang, A. H.Department of Life Science, National Central University, Jung-Li, 320, Taiwan, ROC.Unusual DNA duplex and hairpin motifsNucleic Acids ResNucleic Acids Res2461-7431102003/05/09Base Pair Mismatch/geneticsBase Pairing/geneticsBase SequenceDNA/*chemistry/geneticsHydrogen BondingModels, Molecular*Nucleic Acid ConformationNucleic Acid Heteroduplexes/*chemistry/genetics2003May 151362-4962 (Electronic)12736295http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12736295156048eng(Chou, Chin, and Wang 2003). The scheme of the unusual 3-D structure of AATGG repeats is illustrated on the Figures 8- 10 of Catasti and co-authors  ADDIN EN.CITE Catasti199944417Catasti, P.Chen, X.Mariappan, S. V.Bradbury, E. M.Gupta, G.Life Sciences Division, Los Alamos National Laboratory, NM 87545, USA.DNA repeats in the human genomeGeneticaGenetica15-361061-22000/03/11*Antigens, NuclearBase PairingBase SequenceCentromere/geneticsCpG IslandsDNA/chemistry/*genetics*DNA HelicasesDNA ReplicationDNA-Binding Proteins/genetics*Genome, HumanHumansInsulin/genetics*Interspersed Repetitive SequencesMagnetic Resonance SpectroscopyMethylationModels, MolecularNuclear Proteins/geneticsNucleic Acid ConformationTrinucleotide Repeats19990016-6707 (Print)10710707http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10710707eng(Catasti et al. 1999) while a slightly different variant of spatial organization of the same repeat is illustrated in Figure 1 of Jaishree and Wang  ADDIN EN.CITE Jaishree199411117Jaishree, T. N.Wang, A. H.Biophysics Division, University of Illinois at Urbana-Champaign 61801.Human chromosomal centromere (AATGG)n sequence forms stable structures with unusual base pairsFEBS LettFEBS Lett99-10334711994/06/20Base CompositionBase Sequence*Centromere*Chromosomes, HumanDNA, Satellite/*chemistryHumansMagnetic Resonance SpectroscopyModels, MolecularMolecular Sequence DataNucleic Acid Conformation*Repetitive Sequences, Nucleic Acid1994Jun 200014-5793 (Print)8013671http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=80136710014-5793(94)00516-8 [pii]eng(Jaishree and Wang 1994). These two papers demonstrate that the AATGG strand of the repeat forms stable doubly folded hairpins with Watson-Crick A-T and non-Watson-Crick A-G and G-G base pairs in the stems. The stability of these stems is reached partially due to stacking of the three purines shown by arrows in Figure 1. Moreover, the same authors demonstrated that a greater number of the AATGG pentamers might form higher order structure in which doubly folded hairpins are compactly organized in a helical array of (AATGG)4 units. At the same time, the complementary strand formed by CCATT pentamers is unstable under physiological conditions and likely represents loose structures. Under acidic conditions this CCATT tandem repeat might also form unusual structures known as i-motif with intercalated cytosine bases shown in Figure 2  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Catasti et al. 1999; Nonin-Lecomte and Leroy 2001). Other tandemly organized simple repeats could also have conformations far distinct from Watson-Crick double helices. One of the prominent non-canonical structures, known as G-quadruplex, G-quartet, or G4, is formed by guanine-rich strands of the repeats. Quadruplexes are arranged in four-stranded structures with stands connected to each other via Hoogsteen hydrogen bonding. G-quadruplex has been well-characterized in human telomeric and related sequences with the core repetitive element TTAGGGG and also within promoters and 5-untranslated regions of human genes whose sequences have a loose consensus of G3-5NL1G3-5NL2G3-5NL3G3-5 , where NL1, NL2, and NL3 are loops with the length from 1 to 7 nucleotides and variable nucleotide composition  ADDIN EN.CITE Neidle200911111117Neidle, S.Cancer Research UK Biomolecular Structure Group, The School of Pharmacy, University of London, 29-39 Brunswick Square, London WC1N 1AX, UK. stephen.neidle@pharmacy.ac.ukThe structures of quadruplex nucleic acids and their drug complexesCurr Opin Struct BiolCurr Opin Struct Biol239-501932009/06/03AnimalsBase Sequence*G-QuadruplexesGenomicsHumansLigandsNucleic Acids/*chemistry/genetics/*metabolismPharmaceutical Preparations/*metabolismTelomere/genetics/metabolism2009Jun1879-033X (Electronic)19487118http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=19487118S0959-440X(09)00046-3 [pii] 10.1016/j.sbi.2009.04.001eng(Neidle 2009). There are several alternative conformations of G-quadruplexes due to the organization of the strands relative to each other. Among them are anti-parallel, parallel, and parallel/anti-parallel hybrids  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Oganesian and Bryan 2007; Huppert 2008). One example of a parallel-stranded G4 NMR structure is illustrated in Figure 3. G-quadruplex structure has been demonstrated for several G-rich short tandem repeats. Among them are GGA and CGG triplet repeats ( ADDIN EN.CITE  ADDIN EN.CITE.DATA (Matsugami et al. 2003; Nakagama et al. 2006) and the GGCAG mouse minisatellite Pc-1  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Katahira et al. 1999). Four elements of GGA-repeat can form intramolecular parallel quadruplex, while the neighboring quadruplexes can form a dimer stabilized through the stacking interaction between the heptads of the two quadruplexes  ADDIN EN.CITE Matsugami200324024024017Matsugami, A.Okuizumi, T.Uesugi, S.Katahira, M.Department of Environment and Natural Sciences, Graduate School of Environment and Information Sciences, Yokohama National University, 79-7 Tokiwadai, Hodogaya-ku, Yokohama 240-8501, Japan.Intramolecular higher order packing of parallel quadruplexes comprising a G:G:G:G tetrad and a G(:A):G(:A):G(:A):G heptad of GGA triplet repeat DNAJ Biol ChemJ Biol Chem28147-53278302003/05/16Circular DichroismCrystallography, X-RayDNA/*chemistry/metabolismGuanine/*chemistryIonsMagnetic Resonance SpectroscopyModels, MolecularNucleic Acid ConformationPotassium/metabolism*Repetitive Sequences, Nucleic AcidSodium/metabolismTelomere/ultrastructure*Trinucleotide Repeats2003Jul 250021-9258 (Print)12748183http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1274818310.1074/jbc.M303694200 M303694200 [pii]eng(Matsugami et al. 2003). While G-quadruplexes are formed by a guanine-rich strand, their complementary strand being C-rich may also form a completely different four-stranded structure known as i-motif or intercalated cytosine tetraplex. This structure is only stable under acidic conditions  ADDIN EN.CITE Huppert200817717717717Huppert, J. L.Cavendish Laboratory, University of Cambridge, JJ Thomson Ave, Cambridge, UKCB3 0HE. jlh29@cam.ac.ukFour-stranded nucleic acids: structure, function and targeting of G-quadruplexesChem Soc RevChem Soc Rev1375-843772008/06/24AcridinesBase PairingBinding SitesChemistry, Pharmaceutical/*methods*G-QuadruplexesGene Targeting/*methods*Nucleic Acid ConformationNucleic Acids/*chemistry/genetics/metabolismOxazolesPorphyrinsTelomerase/antagonists & inhibitorsTranscription, Genetic2008Jul0306-0012 (Print)18568163http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1856816310.1039/b702491feng(Huppert 2008). Intercalated cytosines have been found in several unusual conformations, one of which is shown in Figure 2. There are up to ten different non-B DNA conformations associated with the simple repeats listed and well-illustrated by Wells  ADDIN EN.CITE Wells200725125125117Wells, R. D.Center for Genome Research, Institute of Biosciences and Technology, Texas A&M University System Health Science Center, Texas Medical Center, 2121 West Holcombe Boulevard, Houston, TX 77030-3303, USA. rwells@ibt.tamhsc.eduNon-B DNA conformations, mutagenesis and diseaseTrends Biochem SciTrends Biochem Sci271-83262007/05/12Base SequenceDNA/*chemistry/*geneticsDNA, Bacterial/chemistryDNA, Superhelical/chemistryGenetic Diseases, Inborn/geneticsHumansMutagenesis/*genetics*Nucleic Acid ConformationPlasmids/genetics2007Jun0968-0004 (Print)17493823http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=17493823S0968-0004(07)00091-6 [pii] 10.1016/j.tibs.2007.04.003eng(Wells 2007). Among them are slipped structure formed by CNG repeats; triplex DNA formed by purine (R)-rich or pyrimidine (Y)-rich mirror repeats (described below in details); sticky DNA formed by G+A-rich tracts like (GAA)N; and DNA unwinding elements formed by A+T-rich regions. In addition, Wells describes cruciforms created by inverted repeats; and left-handed Z-DNA formed by alternated R/Y bases in (RY)N repeats  ADDIN EN.CITE Wells200725125125117Wells, R. D.Center for Genome Research, Institute of Biosciences and Technology, Texas A&M University System Health Science Center, Texas Medical Center, 2121 West Holcombe Boulevard, Houston, TX 77030-3303, USA. rwells@ibt.tamhsc.eduNon-B DNA conformations, mutagenesis and diseaseTrends Biochem SciTrends Biochem Sci271-83262007/05/12Base SequenceDNA/*chemistry/*geneticsDNA, Bacterial/chemistryDNA, Superhelical/chemistryGenetic Diseases, Inborn/geneticsHumansMutagenesis/*genetics*Nucleic Acid ConformationPlasmids/genetics2007Jun0968-0004 (Print)17493823http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=17493823S0968-0004(07)00091-6 [pii] 10.1016/j.tibs.2007.04.003eng(Wells 2007). There is evidence that CGG triplet repeats could form a non-B-type higher order structure  ADDIN EN.CITE Nakagama200618418418417Nakagama, H.Higuchi, K.Tanaka, E.Tsuchiya, N.Nakashima, K.Katahira, M.Fukuda, H.Biochemistry Division, National Cancer Center Research Institute, 5-1-1 Tsukiji, Tokyo 104-0045, Japan. hnakagam@gan2.res.ncc.go.jpMolecular mechanisms for maintenance of G-rich short tandem repeats capable of adopting G4 DNA structuresMutat ResMutat Res120-315981-22006/03/043T3 CellsAnimalsBase SequenceCircular DichroismDNA/*chemistry*GuanineMiceMolecular WeightNucleic Acid ConformationPolymorphism, GeneticTandem Repeat SequencesTrinucleotide Repeats2006Jun 250027-5107 (Print)16513142http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=16513142S0027-5107(06)00057-1 [pii] 10.1016/j.mrfmmm.2006.01.014eng(Nakagama et al. 2006). This particular CGG repeat is widespread in animal genomes and it expansion inside the first exon of the FMR1 gene that causes Fragile X syndrome  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Mandel 1993; Crawford, Acuna, and Sherman 2001). NMR and X-ray crystallography studies of DNA oligonucleotides strongly suggest that (GA)N and (A)N repeats could form parallel-stranded homeoduplexes  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Kypr, Kejnovska, and Vorlickova 2007; Chakraborty et al. 2009). However it is questionable whether such structures might exist in vivo. Tandem repeats with longer units could also form non-B structures. For instance, the (ACAGGGGTGTGGGG)N insulin minisatellite has a complex loop-folding conformation  ADDIN EN.CITE Catasti199944417Catasti, P.Chen, X.Mariappan, S. V.Bradbury, E. M.Gupta, G.Life Sciences Division, Los Alamos National Laboratory, NM 87545, USA.DNA repeats in the human genomeGeneticaGenetica15-361061-22000/03/11*Antigens, NuclearBase PairingBase SequenceCentromere/geneticsCpG IslandsDNA/chemistry/*genetics*DNA HelicasesDNA ReplicationDNA-Binding Proteins/genetics*Genome, HumanHumansInsulin/genetics*Interspersed Repetitive SequencesMagnetic Resonance SpectroscopyMethylationModels, MolecularNuclear Proteins/geneticsNucleic Acid ConformationTrinucleotide Repeats19990016-6707 (Print)10710707http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10710707eng(Catasti et al. 1999). The listed non-B DNA conformations likely do not exist permanently, but only under specific conditions. Their formation can be facilitated by negative supercoiling during transcription or by binding with transcription factors  ADDIN EN.CITE Mirkin200826626626617Mirkin, S. M.Department of Biology, Tufts University, Medford, MA 02155, USA. Sergei.Mirkin@tufts.eduDiscovery of alternative DNA structures: a heroic decade (1979-1989)Front BiosciFront Biosci1064-71132007/11/06Biochemistry/history/trendsDNA/*chemistryDNA ReplicationDNA Restriction EnzymesDNA, CruciformDNA, Z-FormG-QuadruplexesGenetics/*historyHistory, 20th CenturyHumans*Nucleic Acid ConformationRepetitive Sequences, Nucleic Acid20081093-4715 (Electronic)17981612http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=179816122744 [pii]eng(Mirkin 2008). The very specific pattern of mutagenesis within simple repeats associated with particular bases and particular sites strongly suggests the existence of non-B structures in vivo  ADDIN EN.CITE Wells200725125125117Wells, R. D.Center for Genome Research, Institute of Biosciences and Technology, Texas A&M University System Health Science Center, Texas Medical Center, 2121 West Holcombe Boulevard, Houston, TX 77030-3303, USA. rwells@ibt.tamhsc.eduNon-B DNA conformations, mutagenesis and diseaseTrends Biochem SciTrends Biochem Sci271-83262007/05/12Base SequenceDNA/*chemistry/*geneticsDNA, Bacterial/chemistryDNA, Superhelical/chemistryGenetic Diseases, Inborn/geneticsHumansMutagenesis/*genetics*Nucleic Acid ConformationPlasmids/genetics2007Jun0968-0004 (Print)17493823http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=17493823S0968-0004(07)00091-6 [pii] 10.1016/j.tibs.2007.04.003eng(Wells 2007). Also, several non-B structures have been confirmed in various experiments including in vivo studies  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Wells 2007; Fernando et al. 2009; Kypr et al. 2009). Currently, more than 70 human genetic disorders have been associated with changes in simple repeats  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Lupski 1998; Wells 2007). In summary, simple repeats are abundant in the genomes of diverse animals and plants. In rodents, 2.4% of the euhromatic part of their genome is represented by simple repeats, which is two times bigger than the length of all protein coding sequences  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Gibbs et al. 2004). Additionally, tandemly organized short sequences are abundant and are key components of telomeres and centromeric regions. Many simple repeats, whose total length reaches 20 bases and above, under certain conditions can exist in a variety of non-B DNA conformations in vivo associated with specific genomic functions. Computationally, simple repeats can be detected by the RepeatMasker program  ADDIN EN.CITE Smit AFA52352352312Smit AFA, Hubley R, Green P RepeatMasker Open-3.1.8 http://www.repeatmasker.org(Smit AFA). However, the default parameters of this program could skip recognition of simple repeat loci whose copy numbers are low or where the sequences have accumulated mutations (fuzzy repeats). In this case the best choice is the stand-alone Tandem Repeat Finder with advanced search parameters  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Benson 1999). Genomic Mid-Range Inhomogeneity (MRI): nucleotide compositional extremes and sequence non-randomness In thousands of genomic regions the composition of A, T, C, or G content or different combinations of these bases exist at extremes far different from the average base composition. We call such compositional extremes genomic mid-range inhomogeneity or MRI if they stretch at least 30 base pairs but less than ten thousand base pairs. To characterize genomic MRI patterns, a public computational resource (Genomic MRI) has been created that allows detecting sequence regions with any type of extreme composition  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Bechtel et al. 2008). Using this resource it was demonstrated that various MRI regions occupy up to a quarter of the human genome and their existence is maintained via strong fixation bias  ADDIN EN.CITE Prakash200952652652617<style face="bold" font="Times New Roman" size="100%">Evolution of Genomic Sequence Inhomogeneity at Mid-range Scales </style>BMC GenomicsBMC Genomics5131020091471-216410.1186/1471-2164-10-513(Prakash et al. 2009). Genomic MRI toolkit. For examining mid-range sequence patterns Genomic MRI programs do not characterize particular words but only the overall compositional content of particular base(s) that we refer to as X (X could be a single nucleotide A, G, C, or T or any of their combinations like A+C, or G+T+C, etc.). Genomic MRI allows studying the distribution of X-rich regions in any sequence of interest. These X-rich MRI regions are highly over-represented in mammalian genomes for all kinds X-contexts. For instance, in the human genome G+C-rich sequences with lengths from 100 to 200 nucleotides are 20 times over-represented; A+T-rich sequences in the same length range are about 12 times over-represented; A+G-rich and T+C-rich sequences 10 times; and G+T-rich and A+C-rich sequences up to 6 times over-represented  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Bechtel et al. 2008). In order to measure the abundance of X-rich regions in the sequences under analysis Genomic MRI compares their presence inside a specifically generated random sequence that has the same oligonucleotide distribution as the real one. This evaluation is achieved by the following computational steps. Firstly, the short-range inhomogeneity (SRI) of a given sequence is analyzed by the SRI-analyzer program from the Genomic MRI package to create an oligonucleotide frequency table for each possible 1-9 nucleotide long word. Then, a second program, SRI-generator, creates a random sequence with a short-range inhomogeneity that approximates the oligonucleotide frequency table of the natural sequence. This random sequence is used further for comparison with the natural one. Finally, the third program, MRI-analyzer, scans a sequence under analysis and the random sequence with a window of a specified size and checks whether the nucleotide composition of the sequence in the current window is X-rich or X-poor for a particular chosen combination of nucleotides (X), e.g., A, T, C, G, G+C, A+G, G+T, et cetera. A window is rich for the X-content if its X-composition is above a user-specified threshold-X1, while a window is X-poor if it is below another user-specified threshold-X2. (Note that X-poor regions can be referred to as non-X-rich ones, e.g. G+C-poor are A+T-rich). An example of MRI analyzer graphical output is shown in Figure 4 that illustrates the MRI-patterns for an extra-large human intron of the dystrophin gene from chromosome X. Two scales of MRI regions should be considered. First, regions from 30 to 1000 bp, whose properties have investigated in detail and for which several periodicities have been reported  ADDIN EN.CITE Trifonov1991393917Trifonov, E. N.Department of Structural Biology, Weizmann Institute of Science, Rehovot, Israel.DNA in profileTrends Biochem Sci467-701612Base SequenceDNA/ chemistryMolecular ConformationMolecular Sequence DataStructure-Activity Relationship1991Dec0968-0004 (Print)1781024engIoshikhes1999383817Ioshikhes, I.Trifonov, E. N.Zhang, M. Q.Cold Spring Harbor Laboratory, Cold Spring Harbor, NY 11724, USA.Periodical distribution of transcription factor sites in promoter regions and connection with chromatin structureProc Natl Acad Sci U S A2891-5966AnimalsBinding Sites/geneticsChromatin/ genetics/ultrastructureGenome, HumanHumansPromoter Regions (Genetics)/ geneticsTranscription Factors/ geneticsTranscription, Genetic1999Mar 160027-8424 (Print)10077607engHerzel1999373717Herzel, H.Weiss, O.Trifonov, E. N.Institute for Theoretical Biology, Humboldt University, Invalidenstr. 43, D-10115 Berlin, Germany. h.herzel@biologie.hu-berlin.de10-11 bp periodicities in complete genomes reflect protein structure and DNA foldingBioinformatics187-93153Archaea/geneticsBacteria/geneticsDNA/ chemistry/ geneticsDNA, Archaeal/chemistry/geneticsDNA, Bacterial/chemistry/geneticsDNA, Fungal/chemistry/geneticsDNA, Superhelical/chemistry/geneticsDatabases, FactualFungal Proteins/chemistry/geneticsGenomeGenome, FungalNucleic Acid ConformationNucleosomes/geneticsProtein ConformationRepetitive Sequences, Nucleic AcidSoftwareYeasts/genetics1999Mar1367-4803 (Print)10222405eng(Trifonov 1991; Herzel, Weiss, and Trifonov 1999; Ioshikhes, Trifonov, and Zhang 1999). Second, larger regions from 1 to 10 kb, which are one of the least studied areas in genomic composition and where as yet unknown biological properties may be found. Such subdivisions are important for the proper choice of parameters for the MRI thresholds. For instance, for a 100 nucleotide-long window there are a vast number of regions in mammals where G+C composition is 85% or higher. However, for studying regions with a window-size of around 5 kb, the upper threshold for G+C content should not be more than 65% to find the areas satisfying the criterion. Extended regions with compositional extremes satisfying G+C- or A+T-richness are abundant in vertebrates and can be as long as several million bases (known as genomic isochors). Other composition extremes, such as R-, Y-, G+T-, A+C-richness, that extend over long chromosomal regions are not as abundant as C+G- and A+T- rich genomic areas. Nonetheless, for more than twenty-one hundred human chromosomal regions with lengths exceeding 10 kb we have detected frequencies of more than 60% for G+A-, T+C-, A+C-, or G+T- nucleotides. As for extremes, our computations have shown that there are 22 regions in the human genome where R or Y composition exceeds 70% within a sequence longer than 10 kb. Recently, by studying the distribution of more than four million SNPs in the human genome and by taking into account their frequencies in the population, the influence of mutations on different MRI regions has been examined  ADDIN EN.CITE Prakash200952652652617<style face="bold" font="Times New Roman" size="100%">Evolution of Genomic Sequence Inhomogeneity at Mid-range Scales </style>BMC GenomicsBMC Genomics5131020091471-216410.1186/1471-2164-10-513(Prakash et al. 2009). The authors demonstrated that MRI regions have comparable levels of de novo mutations to the control genomic sequences with average base composition. De novo substitutions rapidly erode MRI regions, bringing their nucleotide composition toward genome-average levels. However, those substitutions that favor the maintenance of MRI properties have a higher chance to spread through the entire population. All in all, the observed strong fixation bias for mutations helps to preserve MRI regions during evolution indicating on their involvement in genomic operations. G+C-rich and A+T-rich MRI regions are associated with several unusual DNA structures We start considering mid-range genomic compositional patterns from the most studied case: G+C-rich and A+T-rich regions. These G+C-rich and A+T-rich regions of various lengths from thirty to several thousand nucleotides are 4-20 times over-represented in the mammalian genomes compared to random expectation  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Bechtel 2008; Bechtel et al. 2008). Among G+C-rich genomic segments, CpG-islands have drawn the most public attention, due to their functional properties and involvement in gene expression regulation  ADDIN EN.CITE Hackenberg200627727727717Hackenberg, M.Previti, C.Luque-Escamilla, P. L.Carpena, P.Martinez-Aroza, J.Oliver, J. L.Dpto. de Genetica, Facultad de Ciencias, Universidad de Granada, Spain. mlhack@gmail.comCpGcluster: a distance-based algorithm for CpG-island detectionBMC BioinformaticsBMC Bioinformatics44672006/10/14*AlgorithmsAnimalsCpG Islands/*geneticsGenome/geneticsHumansMice20061471-2105 (Electronic)17038168http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1703816816171221471-2105-7-446 [pii] 10.1186/1471-2105-7-446eng(Hackenberg et al. 2006). CpG-islands are found in nearly 60% of human genes including almost all of the housekeeping ones  ADDIN EN.CITE Hackenberg200627727727717Hackenberg, M.Previti, C.Luque-Escamilla, P. L.Carpena, P.Martinez-Aroza, J.Oliver, J. L.Dpto. de Genetica, Facultad de Ciencias, Universidad de Granada, Spain. mlhack@gmail.comCpGcluster: a distance-based algorithm for CpG-island detectionBMC BioinformaticsBMC Bioinformatics44672006/10/14*AlgorithmsAnimalsCpG Islands/*geneticsGenome/geneticsHumansMice20061471-2105 (Electronic)17038168http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1703816816171221471-2105-7-446 [pii] 10.1186/1471-2105-7-446eng(Hackenberg et al. 2006). According to two different definitions of these islands, their length must be at least 200 or 500 bp long; G+C content more than 50 or 55%; and the number of CpG dinucleotides in the islands should exceed more than twice their occurrence in other genomic regions [ ADDIN EN.CITE  ADDIN EN.CITE.DATA (Gardiner-Garden and Frommer 1987; Takai and Jones 2003), respectively]. CpG dinucleotides are important sites for cytosine methylation in all vertebrates and some invertebrates and plants. However, inside CpG-islands CpG dinucleotides are predominantly non-methylated  ADDIN EN.CITE Suzuki200828728728717Suzuki, M. M.Bird, A.The Wellcome Trust Centre for Cell Biology, The University of Edinburgh, Michael Swann Building, The King's Buildings, Edinburgh EH9 3JR, UK. a.bird@ed.ac.ukDNA methylation landscapes: provocative insights from epigenomicsNat Rev GenetNat Rev Genet465-76962008/05/09Animals*DNA Methylation*Epigenesis, Genetic*GenomeGenomics/*methodsHumansTranscription, Genetic2008Jun1471-0064 (Electronic)18463664http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=18463664nrg2341 [pii] 10.1038/nrg2341eng(Suzuki and Bird 2008). It has been shown recently that CpG dinucleotide without methylation exhibit structural abnormalities in the DNA helix. Particularly, they are one of the most frequent sites for DNA backbone cleavage by hydroxyl radicals  ADDIN EN.CITE Greenbaum2007262617Greenbaum, J. A.Parker, S. C.Tullius, T. D.Program in Bioinformatics, Boston University, Boston, Massachusetts 02215, USA.Detection of DNA structural motifs in functional genomic elementsGenome Res940-6176Chromatin/ geneticsComputational BiologyDatabases, GeneticGenome, HumanHumansRegulatory Elements, TranscriptionalSequence Analysis, DNA2007Jun1088-9051 (Print)17568009engGreenbaum2007252517Greenbaum, J. A.Pang, B.Tullius, T. D.Program in Bioinformatics, Boston University, Boston, Massachusetts 02215, USA.Construction of a genome-scale structural map at single-nucleotide resolutionGenome Res947-53176AlgorithmsChromosome MappingDatabases, GeneticGene LibraryGenome, HumanHumansNucleotides/geneticsPredictive Value of TestsSequence Analysis, DNA2007Jun1088-9051 (Print)17568010eng(Greenbaum, Pang, and Tullius 2007; Greenbaum, Parker, and Tullius 2007) and during the sonication of double-stranded DNA  ADDIN EN.CITE Grokhovsky2008292917Grokhovsky, S L, Il'icheva, I A,Nechipurenko, D Yu.,Panchenko, L A,Polozov, R L,Nechipurenko, Yu DHeterogeneity of DNA local structure and dynamics: ultrasound studies. Biofizika417-425532008(Grokhovsky et al. 2008). The crucial involvement of cytosine methylation in the regulation of gene expression is well described in a number of reviews including some recent ones  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Prokhortchouk and Defossez 2008; Suzuki and Bird 2008; Illingworth and Bird 2009). Thus, here we concentrate on the other physicochemical properties of G+C-rich and A+T-rich regions. It is well known that A-form of DNA helix exists in high salt concentrations and in ethanol-containing solutions. However, G+C-rich regions may be present in A-form DNA even in aqueous solutions  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Warne and deHaseth 1993; Stefl et al. 2001; Kypr et al. 2009). A special form of DNA which is an intermediate between A- and B-forms, has been characterized in G+C-rich sequences with methylated cytosines  ADDIN EN.CITE Vargason200029529529517Vargason, J. M.Eichman, B. F.Ho, P. S.Department of Biochemistry and Biophysics, ALS 2011, Oregon State University, Corvallis, Oregon 97331-7305, USA.The extended and eccentric E-DNA structure induced by cytosine methylation or brominationNat Struct BiolNat Struct Biol758-61792000/08/31Base SequenceBromine/*metabolismCpG Islands/geneticsCrystallizationCrystallography, X-RayCytosine/*metabolismDNA/*chemistry/genetics/*metabolism*DNA MethylationModels, Molecular*Nucleic Acid ConformationSolventsWater/metabolism2000Sep1072-8368 (Print)10966645http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1096664510.1038/78985eng(Vargason, Eichman, and Ho 2000). In addition, short (CpG)n repeats could adopt Z-DNA as recently has been reviewed by P.S. Ho  ADDIN EN.CITE Ho200930130130117Ho, P. S.Methods to study nucleic acid structureMethodsMethods1414732009/02/14Chemistry, Analytical/*methodsGenetic TechniquesMicroscopy/methods*Nucleic Acid ConformationNucleic Acids/*chemistry2009Mar1095-9130 (Electronic)19215858http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=19215858S1046-2023(09)00004-8 [pii] 10.1016/j.ymeth.2009.01.003eng(Ho 2009). This Z-DNA is proposed to serve as transcriptional co-activator  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Liu et al. 2001). A+T-rich regions, on the other hand, are also associated with special DNA conformations. Some of these sequences with specific distributions of A and T bases form an unusual structure known as the DNA unwinding element  ADDIN EN.CITE Kowalski198831331331317Kowalski, D.Natale, D. A.Eddy, M. J.Molecular and Cellular Biology Department, Roswell Park Memorial Institute, Buffalo, NY 14263.Stable DNA unwinding, not "breathing," accounts for single-strand-specific nuclease hypersensitivity of specific A+T-rich sequencesProc Natl Acad Sci U S AProc Natl Acad Sci U S A9464-885241988/12/01Base SequenceBinding SitesDNA, Superhelical/*metabolismEndonucleases/*metabolismMolecular Sequence Data*Poly A*Poly T*PolydeoxyribonucleotidesSingle-Strand Specific DNA and RNA EndonucleasesStructure-Activity Relationship1988Dec0027-8424 (Print)2849106http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=2849106282773eng(Kowalski, Natale, and Eddy 1988). These elements are often associated with the origins of replication in eukaryotes and prokaryotes  ADDIN EN.CITE Umek198931531531517Umek, R. M.Linskens, M. H.Kowalski, D.Huberman, J. A.Department of Molecular and Cellular Biology, Roswell Park Memorial Institute, Buffalo, NY 14263.New beginnings in studies of eukaryotic DNA replication originsBiochim Biophys ActaBiochim Biophys Acta1-14100711989/01/23Base SequenceCells/*ultrastructure*DnaDNA-Binding Proteins/physiologyEukaryotic Cells/*ultrastructureGene Expression RegulationModels, Genetic*RepliconSaccharomyces cerevisiae/genetics1989Jan 230006-3002 (Print)2642383http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=2642383eng(Umek et al. 1989). There are several A+T-rich simple repeats widespread in eukaryotes. Among them, (AT)n is one of the most common in animals. X-ray and NMR studies of the DNA oligomer d(ATATAT) have shown that in addition to B-DNA it could form an anti-parallel double helical duplex in which the base pairing is of the Hoogsteen type  ADDIN EN.CITE Abrescia200431631631617Abrescia, N. G.Gonzalez, C.Gouyette, C.Subirana, J. A.Departament d'Enginyeria Quimica, Universitat Politecnica de Catalunya, Av. Diagonal 647, E-08028 Barcelona, Spain. nicola@strubi.ox.ac.ukX-ray and NMR studies of the DNA oligomer d(ATATAT): Hoogsteen base pairing in duplex DNABiochemistryBiochemistry4092-10043142004/04/07*Base PairingCrystallizationCrystallography, X-RayDNA/*chemistryNuclear Magnetic Resonance, BiomolecularNucleic Acid ConformationNucleic Acid Heteroduplexes/*chemistryOligonucleotides/*chemistrySolutions2004Apr 130006-2960 (Print)15065851http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1506585110.1021/bi0355140eng(Abrescia et al. 2004). The adenines in this duplex are flipped over making the minor groove narrow and hydrophobic. This structure is very similar to the standard B-form helix with about 10 base pairs per turn. Theoretical analysis has demonstrated that energies of the Hoogsteen form and B-form of DNA are practically identical  ADDIN EN.CITE Cubero200331831831817Cubero, E.Abrescia, N. G.Subirana, J. A.Luque, F. J.Orozco, M.Molecular Modeling & Bioinformatic Unit, Institut de Recerca Biomedica, Parc Cientific de Barcelona, Josep Samitier 1-5, Barcelona 08028, Spain.Theoretical study of a new DNA structure: the antiparallel Hoogsteen duplexJ Am Chem SocJ Am Chem Soc14603-12125472003/11/20DNA/*chemistryModels, MolecularNucleic Acid ConformationPoly dA-dT/*chemistrySolutionsThermodynamicsWater/chemistry2003Nov 260002-7863 (Print)14624611http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1462461110.1021/ja035918feng(Cubero et al. 2003). Most recently, Chakraborty and co-authors demonstrated that poly-dA oligonucleotides (dA15) under acidic pH conditions could allow the formation of a double-helical parallel-stranded duplex held together by reversed Hoogsteen type AH+-H+A base pairs  ADDIN EN.CITE Chakraborty200925625625617Chakraborty, S.Sharma, S.Maiti, P. K.Krishnan, Y.National Centre for Biological Sciences, TIFR, GKVK, Bangalore, India.The poly dA helix: a new structural motif for high performance DNA-based molecular switchesNucleic Acids ResNucleic Acids Res2810-73792009/03/13Circular DichroismComputer SimulationDNA/*chemistryDimerizationElectrophoresis, Polyacrylamide GelHydrogen-Ion ConcentrationModels, MolecularNuclear Magnetic Resonance, BiomolecularNucleic Acid ConformationNucleic Acid DenaturationPoly A/*chemistrySodium Chloride/chemistrySpectrometry, FluorescenceTemperature2009May1362-4962 (Electronic)19279188http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=192791882685084gkp133 [pii] 10.1093/nar/gkp133eng(Chakraborty et al. 2009). A+T-rich regions presumably have several important cellular functions. First, the most indicative compositional characteristic of scaffold/matrix-attached regions is that they are A+T-rich  ADDIN EN.CITE Liebich200232032032017Liebich, I.Bode, J.Reuter, I.Wingender, E.Research Group Bioinformatics, Gesellschaft fur Biotechnologische Forschung mbH, Mascheroder Weg 1, D-38124 Braunschweig, Germany. ili@gbf.deEvaluation of sequence motifs found in scaffold/matrix-attached regions (S/MARs)Nucleic Acids ResNucleic Acids Res3433-4230152002/07/26AT Rich SequenceAnimalsBinding SitesConsensus SequenceDatabases, Nucleic AcidExonsNuclear Matrix/geneticsRegulatory Sequences, Nucleic AcidSequence Analysis, DNA/*methodsTranscription Factors/metabolism2002Aug 11362-4962 (Electronic)12140328http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12140328137072eng(Liebich et al. 2002). Second, centromere DNA of diverse animals, plants and fungi always contain A+T-rich regions  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Choo 1997; Abrescia et al. 2004) R-rich/Y-rich MRI regions are associated with H-DNA triplex All combinations of nucleotide pairs except G+C and A+T have strand asymmetry. For example, if one strand is enriched by purines (R) the complementary strand is enriched by pyrimidines (Y). Therefore, R- and Y-rich sequences and also T+G-rich and A+C-rich ones are physically the same loci, yet representing complementary strands. From here on we will consider them together and refer to as R/Y-rich and T+G/A+C-rich respectively. Since 1957 it has been shown that complementary DNA strands, one of which is R-rich and another Y-rich, can form three-stranded helical structures or triplexes  ADDIN EN.CITE Felsenfeld195733433433417Felsenfeld, G.Rich, A.Studies on the formation of two- and three-stranded polyribonucleotidesBiochim Biophys ActaBiochim Biophys Acta457-682631957/12/01*Nucleic Acids1957Dec0006-3002 (Print)13499402http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=13499402eng(Felsenfeld and Rich 1957). Intramolecular triplexes, known also as H-DNA, materialize under certain conditions, like supercoiling, when half of the DNA duplex may dissociate into single strands and one of the stand-alone strands can interact via Hoogsteen base pairing with the remaining Watson-Crick DNA duplex along its major groove forming a triplex structure. The remaining stand-alone strand stays unpaired. An example of a DNA triplex is shown in Figure 5. There are four kinds of H-DNA depending on strand type and orientation  ADDIN EN.CITE Jain200833533533517Jain, A.Wang, G.Vasquez, K. M.Department of Carcinogenesis, University of Texas, M.D. Anderson Cancer Center, Science Park--Research Division, 1808 Park Road 1-C, P.O. Box 389, Smithville, TX 78957, USA.DNA triple helices: biological consequences and therapeutic potentialBiochimieBiochimie1117-309082008/03/12AnimalsAntineoplastic Agents/metabolism/pharmacologyBase SequenceDNA/chemistry/*genetics/*metabolism/*therapeutic useGenome/geneticsHumansNucleic Acid ConformationOligonucleotides/metabolismTranscription, Genetic2008Aug0300-9084 (Print)18331847http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=183318472586808S0300-9084(08)00045-X [pii] 10.1016/j.biochi.2008.02.011eng(Jain, Wang, and Vasquez 2008). One type of H-DNA forms under acidic conditions when the stand-alone Y-rich strand interacts with the R-rich strand of the remaining duplex. Particularly, thymines of the stand-alone strand interact with adenosines of the A-T Watson-Crick pairs of the duplex via Hoogsteen hydrogen bonding, while cytosines of the stand-alone strand interact with guanines of G-C Watson-Crick pairs. Due to this base match requirement for the assembly of this kind of triplex, the sequences of Y-rich stand-alone strand and the Y-rich strand in the duplex should have sequence mirror symmetry. (Here is an example of two sequences with mirror symmetry: 5-TAGTTCC-3 and 5-CCTTGAT-3.) In many R/Y-rich regions of the genomes such mirror symmetry has been observed. For example, a 2.5 kb R-rich sequence of the 21st intron of the human PKD1 gene has 23 mirror repeats that form H-DNA  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Van Raay et al. 1996; Blaszak et al. 1999). Another kind of intramolecular triplex, can be formed at neutral pH and requires bivalent cations for stability. It is formed by the interaction of R-rich stand-alone strand with the remaining duplex via Hoogsteen bonding. It does not require strong mirror symmetry within its sequences, since the adenines of the stand-alone R-rich strand could interact with the A-T pair of the duplex or with the G-C pair  ADDIN EN.CITE Malkov199333933933917Malkov, V. A.Voloshin, O. N.Veselkov, A. G.Rostapshov, V. M.Jansen, I.Soyfer, V. N.Frank-Kamenetskii, M. D.Institute of Molecular Genetics, Russian Academy of Science, Moscow.Protonated pyrimidine-purine-purine triplexNucleic Acids ResNucleic Acids Res105-112111993/01/11Base SequenceDNA/*chemistryDensitometryHydrogen BondingHydrogen-Ion ConcentrationMolecular Sequence DataNucleic Acid ConformationProtonsPurines/*chemistryPyrimidines/*chemistrySulfuric Acid Esters/chemistry1993Jan 110305-1048 (Print)8382787http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8382787309071eng(Malkov et al. 1993). There are several documented functions of H-DNA. It is well established that H-DNA could exist in vivo under certain conditions. Various experimental methods for the characterization of H-DNA have been reviewed recently  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Jain, Wang, and Vasquez 2008; Wang, Zhao, and Vasquez 2009). Single stranded DNA not participating in the triplex is accessible to S1-nuclease cleavage. Eukaryotic genomes contain many S1-nuclease sensitive sites within runs of homo-purine sequences. These segments of single-stranded DNA are frequently involved in the recombination of homologous DNA and thus are sites for genetic instability. Different schemes of recombination involving H-DNA have been described by Jain and others  ADDIN EN.CITE Jain200833533533517Jain, A.Wang, G.Vasquez, K. M.Department of Carcinogenesis, University of Texas, M.D. Anderson Cancer Center, Science Park--Research Division, 1808 Park Road 1-C, P.O. Box 389, Smithville, TX 78957, USA.DNA triple helices: biological consequences and therapeutic potentialBiochimieBiochimie1117-309082008/03/12AnimalsAntineoplastic Agents/metabolism/pharmacologyBase SequenceDNA/chemistry/*genetics/*metabolism/*therapeutic useGenome/geneticsHumansNucleic Acid ConformationOligonucleotides/metabolismTranscription, Genetic2008Aug0300-9084 (Print)18331847http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=183318472586808S0300-9084(08)00045-X [pii] 10.1016/j.biochi.2008.02.011eng(Jain, Wang, and Vasquez 2008). Bacolla with co-authors characterized nearly three thousand homo-purine tracks in the human genome longer than 100 nucleotides  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Bacolla et al. 2006). They supported evidences for these tracks in promoting recombination and association with higher rates of mutations. In addition, stable H-DNA structures are able to block transcription and replication. Jain and co-authors surveyed the evidences for how H-DNA influences the activity of DNA and RNA polymerases. Finally, Goni and others  ADDIN EN.CITE Goni200634834834817Goni, J. R.Vaquerizas, J. M.Dopazo, J.Orozco, M.Molecular Modeling and Bioinformatics Unit, Institut de Recerca Biomedica, Parc Cientific de Barcelona, Josep Samitier 1-5, Barcelona 08028, Spain. rgoni@mmb.pcb.ub.esExploring the reasons for the large density of triplex-forming oligonucleotide target sequences in the human regulatory regionsBMC GenomicsBMC Genomics6372006/03/29AnimalsBase SequenceComputational BiologyDNA/chemistry/*genetics/*metabolismGenome, Human/geneticsHumansMiceNucleic Acid ConformationNucleotides/geneticsOligonucleotides/chemistry/*genetics/*metabolismPromoter Regions, Genetic/geneticsRegulatory Sequences, Nucleic Acid/*geneticsTranscription Factors/metabolism20061471-2164 (Electronic)16566817http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1656681714358861471-2164-7-63 [pii] 10.1186/1471-2164-7-63eng(Goni et al. 2006) performed a large-scale bioinformatic analysis of the distribution of short R-rich sequences in the human genome. They demonstrated that short R-rich sequences are several times more abundant in the downstream promoter regions compared to other regions and to random expectation models. These short R-rich sequences hold evolutionary conservation between human and mouse, yet; likely they are not direct targets for transcription factors. Goni and coauthors have suggested that these sequences act as pacing fragments in promoter regions and help in the correct positioning of transcription factors. DNA and RNA properties of GT-rich/AC-rich MRI regions Recall that the complementary strands of G+T-rich regions are naturally A+C-rich regions. They co-exist with each other and we consider them interchangeably with respect to their description in the literature. According to nucleic acid nomenclature G or T nucleotides are also known as Keto or K while A or C are known as aMino or M  ADDIN EN.CITE Moss52452452412G. P. MossDepartment of Chemistry, Queen Mary University of London, Mile End Road, London, E1 4NS, UKNomenclature for Incompletely Specified Bases in Nucleic Acid Sequenceshttp://www.chem.qmul.ac.uk/iubmb/misc/naseq.html(Moss). Thus, sometimes these regions are referred to as K.M-tracks or motifs  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Yagil 2004). Bechtel and co-authors demonstrated that G+T-regions are about 5 times more abundant in the mammalian genomes compared to random expectation  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Bechtel et al. 2008). Moreover, these regions practically do not intersect with interspersed DNA repeats at all. In 2004 Yagil demonstrated that K.M motifs are significantly over-represented in the genomes of diverse animals, plants, and fungi. Specifically, K.M motifs are predominant in the D. melanogaster genome, where they outnumber other motifs such as R/Y-rich motifs  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Yagil 2004). Despite their abundance, G+T-rich motifs are much less investigated than other regions with extremes in base compositions. Possible functions that could be associated to G+T-rich regions are the following. Firstly, (CA)N simple repeats are one of the most profuse tandem repeats in mammalian genomes  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Waterston et al. 2002). They also should be considered as alternating R/Y sequence, and, due to this property, associated with a Z-DNA conformation  ADDIN EN.CITE Vogt198836036036017Vogt, N.Rousseau, N.Leng, M.Malfoy, B.Centre de Biophysique Moleculaire, Centre National de la Recherche Scientifique, Orleans, France.A study of the B-Z transition of the AC-rich region of the repeat unit of a satellite DNA from Cebus by means of chemical probesJ Biol ChemJ Biol Chem11826-32263241988/08/25AnimalsBase CompositionBase SequenceCebusChemical PhenomenaChemistry*DNA, SatelliteDiethyl PyrocarbonateHydroxylamineHydroxylaminesImmunoassayMolecular Sequence Data*Nucleic Acid ConformationOsmium TetroxideRepetitive Sequences, Nucleic AcidSulfuric Acid Esters1988Aug 250021-9258 (Print)3214490http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=3214490eng(Vogt et al. 1988), which is considered in the next section. Second, C-rich regions, which could be a component of CA-rich regions, are capable of forming four-stranded intercalated molecules  ADDIN EN.CITE Berger199636336336317Berger, I.Egli, M.Rich, A.Department of Biology, Massachusetts Institute of Technology, Cambridge 02139, USA.Inter-strand C-H...O hydrogen bonds stabilizing four-stranded intercalated molecules: stereoelectronic effects of O4' in cytosine-rich DNAProc Natl Acad Sci U S AProc Natl Acad Sci U S A12116-2193221996/10/29Base CompositionCrystallography, X-Ray*CytosineDNA/*chemistryHydrogen BondingMagnetic Resonance SpectroscopyModels, MolecularNucleic Acid ConformationStereoisomerism1996Oct 290027-8424 (Print)8901542http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=890154237952eng(Berger, Egli, and Rich 1996). We mentioned such structures (i-motifs) above in the Simple Repeat section and present an example of it in Figure 2. Third, telomeres of various eukaryotic species are represented by G+T-rich regions which form G-quadruplexes (see above). Fourth, short G+T-rich regions could represent transcription factor binding sites such as for factor Sp1  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Wang et al. 2009). Intriguingly, G+T-rich oligonucleotides possess antiviral activities. For example, T2(G4T2)3 sequence are virucidal against herpes simplex virus  ADDIN EN.CITE Shogan200643343343317Shogan, B.Kruse, L.Mulamba, G. B.Hu, A.Coen, D. M.Dept. of Biological Chemistry and Molecular Pharmacology, Harvard Medical School, Boston, MA 02115, USA.Virucidal activity of a GT-rich oligonucleotide against herpes simplex virus mediated by glycoprotein BJ VirolJ Virol4740-780102006/04/28AnimalsAntiviral Agents/metabolism/*pharmacologyCercopithecus aethiops*Dinucleotide RepeatsDrug Resistance, Viral/geneticsGuanine/metabolismMutationOligonucleotides/*metabolism/*pharmacologySimplexvirus/*drug effects/*genetics/growth & developmentThionucleotides/metabolism/pharmacologyThymine/metabolismTime FactorsVero CellsViral Envelope Proteins/genetics/*physiology2006May0022-538X (Print)16641267http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=16641267147205380/10/4740 [pii] 10.1128/JVI.80.10.4740-4747.2006eng(Shogan et al. 2006). At the RNA level, C+A-rich sequences within intronic segments could regulate alternative splicing by being binding sites for the hnRNP L protein  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Hui et al. 2005). The presence of C+A-rich sequences at the 3-UTR of mRNA could regulate gene expression at the level of translation  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Hamilton et al. 2008). The distribution of C+A-rich sequences enriched by (CA)N imperfect repeats is highly skewed towards telomeres, and minisatellites can usually be found in the vicinity as well  ADDIN EN.CITE Giraudeau199944044044017Giraudeau, F.Petit, E.Avet-Loiseau, H.Hauck, Y.Vergnaud, G.Amarger, V.Institut de Genetique et Microbiologie, Universite Paris-Sud, 91405 Orsay CEDEX, France.Finding new human minisatellite sequences in the vicinity of long CA-rich sequencesGenome ResGenome Res647-53971999/07/21Bacteriophage P1/geneticsChromosome MappingCosmidsDNA/chemistry/geneticsDNA ProbesDatabases, FactualDinucleotide Repeats/*geneticsGenomic LibraryHumansIn Situ Hybridization, FluorescenceMinisatellite Repeats/*geneticsMolecular Sequence DataSequence Analysis, DNA1999Jul1088-9051 (Print)10413403http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10413403310796eng(Giraudeau et al. 1999). Despite the listed properties associated with G+T-rich regions, they seem significantly under-investigated and may yet reveal unknown important functional properties in the near future. Alternated R/Y MRI regions adopt Z-DNA conformation Left-handed anti-parallel Z-DNA double helix conformation has been first characterized in 1979 by Wang and co-authors for (GC)3 repeats  ADDIN EN.CITE Wang197944144144117Wang, A. H.Quigley, G. J.Kolpak, F. J.Crawford, J. L.van Boom, J. H.van der Marel, G.Rich, A.Molecular structure of a left-handed double helical DNA fragment at atomic resolutionNatureNature680-628257401979/12/13Base SequenceCrystallization*DnaHydrogen BondingNucleic Acid ConformationPolydeoxyribonucleotidesStructure-Activity RelationshipX-Ray Diffraction1979Dec 130028-0836 (Print)514347http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=514347eng(Wang et al. 1979). Detailed Z-DNA structure has been considered elsewhere  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Rich and Zhang 2003; Ho 2009). This particular conformation is characterized by rotation of R bases that adopt syn form and stack over the deoxyribose ring, while Y bases do not adopt unfavourable syn form  ADDIN EN.CITE Ho200930130130117Ho, P. S.Methods to study nucleic acid structureMethodsMethods1414732009/02/14Chemistry, Analytical/*methodsGenetic TechniquesMicroscopy/methods*Nucleic Acid ConformationNucleic Acids/*chemistry2009Mar1095-9130 (Electronic)19215858http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=19215858S1046-2023(09)00004-8 [pii] 10.1016/j.ymeth.2009.01.003eng(Ho 2009). Thus, Z-DNA, which is characterized by alternating pattern of anti-syn conformations, is formed by alternated R/Y sequences  ADDIN EN.CITE Johnston199244744744717Johnston, B. H.Cell and Molecular Biology Laboratory, SRI International, Menlo Park, California 94025.Generation and detection of Z-DNAMethods EnzymolMethods Enzymol127-582111992/01/01AntibodiesBase SequenceDNA/analysis/*chemistryDNA, Superhelical/chemistryDNA-Binding Proteins/chemistryMolecular ProbesMolecular Sequence Data*Nucleic Acid ConformationSpectrum Analysis19920076-6879 (Print)1406305http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1406305eng(Johnston 1992). The latest version of the Genomic MRI package has a new feature allowing the detection of excesses and shortages of alternating bases including R/Y patterns. It reveals that in mammalian genomes there is more than 40 times the overabundance of alternating R/Y stretching over 50-100 bp genomic segments, where RY plus YR comprise more than 80% of all dinucleotides. A considerable portion of these alternating R/Y patterns are represented by short (GC)n, (AC)n, (AT)n, and (TG)n repeats that can alternate with each other and be accompanied by alternating R/Y bases without strong periodic sequence pattern. For example, here is a sequence of a 50 bp segment from the third intron of human heparanase-2 gene highly enriched with alternated R and Y bases: 52 AAATGGATGTGTGTATATATATGAAGTCGATACACACACATATACACATA32 . We showed that such alternating R/Y sequences are plentiful throughout the mammalian genomes either inside introns or within intergenic regions. In 1986 Ho and others developed a ZHUNT program for detection of genomic sequences with high propensity to form Z-DNA  ADDIN EN.CITE Ho198644844844817Ho, P. S.Ellison, M. J.Quigley, G. J.Rich, A.A computer aided thermodynamic approach for predicting the formation of Z-DNA in naturally occurring sequencesEMBO JEMBO J2737-445101986/10/01Base Sequence*Dna*Nucleic Acid ConformationSoftwareStructure-Activity RelationshipThermodynamics1986Oct0261-4189 (Print)3780676http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=37806761167176eng(Ho et al. 1986). They found a high concentration of these sequences near the transcription start sites  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Schroth, Chou, and Ho 1992; Rich and Zhang 2003). Most recently, human genomic Z-DNA segments have been detected experimentally using a Z-DNA binding protein domain as a probe  ADDIN EN.CITE Li200946146146117Li, H.Xiao, J.Li, J.Lu, L.Feng, S.Droge, P.Division of Genomics and Genetics, School of Biological Sciences, Nanyang Technological University, 60 Nanyang Drive, Singapore.Human genomic Z-DNA segments probed by the Z alpha domain of ADAR1Nucleic Acids ResNucleic Acids Res2737-463782009/03/12Adenosine Deaminase/*chemistryBinding SitesCell Line, TumorDNA, Z-Form/*analysis/chemistry*Genome, HumanHumansMolecular Probes/chemistryPolymorphism, Single NucleotideProtein Structure, TertiarySequence Analysis, DNASoftware2009May1362-4962 (Electronic)19276205http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=192762052677879gkp124 [pii] 10.1093/nar/gkp124eng(Li et al. 2009). These authors found an abundance of Z-DNA hotspots located in centromeres of 13 human chromosomes. Z-DNA-forming sequences induce high levels of genetic instability in both mammalian and bacterial cells. These sequences could be causative factors for gene translocations found in leukemias and lymphomas  ADDIN EN.CITE Wang200646446446417Wang, G.Christensen, L. A.Vasquez, K. M.Department of Carcinogenesis, University of Texas M. D. Anderson Cancer Center, Science Park-Research Division, 1808 Park Road 1-C, Smithville, TX 78957, USA.Z-DNA-forming sequences generate large-scale deletions in mammalian cellsProc Natl Acad Sci U S AProc Natl Acad Sci U S A2677-8210382006/02/14AnimalsCOS CellsCell Extracts/chemistryCercopithecus aethiopsDNA/analysis*DNA DamageDNA Replication/drug effects/geneticsDNA, Z-Form/genetics/pharmacology/*physiologyEscherichia coli/genetics*Genomic InstabilityHela CellsHumansMutationPlasmids/genetics*Sequence DeletionTranscription, Genetic2006Feb 210027-8424 (Print)16473937http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1647393714138240511084103 [pii] 10.1073/pnas.0511084103eng(Wang, Christensen, and Vasquez 2006). The discovery of certain classes of proteins bound to Z-DNA with high affinity and specificity indicated to a biological role of this structure. Yet, it is a common view that Z-DNA is an unstable conformation that is formed and disappears during particular physiological activities such as transcription  ADDIN EN.CITE Rich200344444444417Rich, A.Zhang, S.Department of Biology, Massachusetts Institute of Technology, 77 Massachusetts Avenue, Room 68-233, Cambridge, Massachusetts 02139, USA.Timeline: Z-DNA: the long road to biological functionNat Rev GenetNat Rev Genet566-72472003/07/03AnimalsDNA, Superhelical/history/*physiologyDNA-Binding Proteins/history/*physiologyHistory, 20th CenturyHistory, 21st CenturyHumansNucleic Acid Conformation2003Jul1471-0056 (Print)12838348http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1283834810.1038/nrg1115 nrg1115 [pii]eng(Rich and Zhang 2003). Weak periodicities and loose patterns In addition to MRI patterns, there are several weak genomic periodicities and specific signals at the mid-range scale. Many of them are described in The codes of life  ADDIN EN.CITE Barbieri200850450450428Marcello BarbieriThe Codes of Life. The rules of MacroevolutionBiosemionticsThe rules of MacroevolutionThe Codes of Life1first20082008Springer978-1-40206339-8(Barbieri 2008). Wherein, Trifonov reviewed different codes that exist in the genomes at DNA, RNA, and protein levels. He emphasized a special property of genomic sequences to make superposition (overlapping) of the codes they carry. The overlapping is possibly due to degeneracy of the codes and might be useful for organism survivability  ADDIN EN.CITE Peleg200446546546517Peleg, O.Kirzhner, V.Trifonov, E.Bolshoy, A.Genome Diversity Center, Institute of Evolution, Haifa University Mt Carmel, Haifa 31905, Israel. peleg@research.haifa.ac.ilOverlapping messages and survivabilityJ Mol EvolJ Mol Evol520-75942005/01/11AnimalsBase SequenceCodon/*genetics*Evolution, MolecularGenome, BacterialGenome, ViralHumansLikelihood Functions*Models, GeneticMolecular Sequence DataMutationSequence Alignment2004Oct0022-2844 (Print)15638463http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1563846310.1007/s00239-004-2644-5eng(Peleg et al. 2004). Here we consider three types of such patterns in eukaryotic genomes. Chromatin periodicities There exists a non-random positioning of nucleosomes along genomic DNA of eukaryotes  ADDIN EN.CITE Salih200747047047017Salih, F.Salih, B.Trifonov, E. N.Genome Diversity Center, Institute of Evolution, University of Haifa, Mount Carmel, Haifa 31905, Israel.Sequence-directed mapping of nucleosome positionsJ Biomol Struct DynJ Biomol Struct Dyn489-932452007/02/23Base Sequence*DnaDinucleotide Repeats/*geneticsMolecular Sequence DataNucleosomes/chemistry/*geneticsRepetitive Sequences, Nucleic Acid/*geneticsSequence Analysis, DNA2007Apr0739-1102 (Print)17313194http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=17313194d=3028&c=4224&p=15650&do=detail [pii]eng(Salih, Salih, and Trifonov 2007). Nucleosome binding preferences are achieved via sequence-dependent deformational anisotropy of DNA  ADDIN EN.CITE Barbieri200850450450428Marcello BarbieriThe Codes of Life. The rules of MacroevolutionBiosemionticsThe rules of MacroevolutionThe Codes of Life1first20082008Springer978-1-40206339-8(Barbieri 2008). On average, one nucleosome occupies 200 bp including 145 nucleotides that contact its core particle while the rest corresponds to linkers between nucleosomes. Due to this specificity in nucleosome positioning Trifonov and co-authors described sequence features that are repeated with 200- and 400-base periodicities  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Trifonov 1998; Cohanim, Kashi, and Trifonov 2006). Periodicities in protein-coding sequences There are well-known short-range periodicities in coding sequences that exist due to non-symmetry in the genetic code, non-random amino acid appearance and association of neighbouring amino acids within protein sequences, and also regularities in codon bias and context-dependent codon bias  ADDIN EN.CITE Fedorov200247347347317Fedorov, A.Saxonov, S.Gilbert, W.Department of Molecular and Cellular Biology, Harvard University, 16 Divinity Avenue, Cambridge, MA 02138, USA. afedorov@fas.harvard.eduRegularities of context-dependent codon bias in eukaryotic genesNucleic Acids ResNucleic Acids Res1192-73052002/02/28AnimalsArabidopsis/geneticsCaenorhabditis elegans/geneticsCodon/*geneticsDrosophila melanogaster/geneticsEukaryotic Cells/metabolismGenesGenomeGenome, HumanHumansModels, StatisticalNucleotides/genetics2002Mar 11362-4962 (Electronic)11861911http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11861911101244eng(Fedorov, Saxonov, and Gilbert 2002). In addition, there exist longer periodicities in the coding sequences that correspond to modular organization of globular proteins. They extend over 20-30 codons and represent initial protein folding modules  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Aharonovsky and Trifonov 2005; Barbieri 2008). Transcription-associated mutational asymmetry in mammals In 2003 Green et al. demonstrated that the transcribed strands of mammalian DNA have an excess of G+T over A+C due to the difference of particular mutation frequencies  ADDIN EN.CITE Green200347647647617Green, P.Ewing, B.Miller, W.Thomas, P. J.Green, E. D.Howard Hughes Medical Institute and Department of Genome Sciences, University of Washington, Seattle, Washington 98195, USA. phg@u.washington.eduTranscription-associated mutational asymmetry in mammalian evolutionNat GenetNat Genet514-73342003/03/04AnimalsCell LineageChromosomes, Human, Pair 22CpG Islands*DNA Mutational AnalysisDatabases as TopicEvolutionHumansModels, GeneticPapioRNA, Messenger/metabolismSequence Analysis, DNA*Transcription, Genetic2003Apr1061-4036 (Print)12612582http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1261258210.1038/ng1103 ng1103 [pii]<language>eng</language></record></Cite></EndNote>(Green et al. 2003). Specifically, the A!G transition occurs at a 28% higher rate than the complementary transition T!C on the transcribed strand in most human genes. This transcription-associated mutational bias exists for both the exonic and intronic parts of genes. Thus, if we look at the nucleotide frequencies in the combined sequences of all human introns (T=30.7%; A=28.0%; G=21.1%; C=20.2%) there is a 3.6% excess of G+T over A+C. [These calculations were obtained on our non-redundant set of 11,315 human genes containing 96,931 introns  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Bechtel et al. 2008). For each intron we removed the first 10 and the last 30 bases]. We detected the same preference of G+T over A+C in introns of other mammals, a smaller preference for sea urchin (2.2%), and the highest preference in Arabidopsis (11.8%). For the mouse-ear cress, the nucleotide bias in introns is mainly due to significant excess of T (39.6%) over A (28.2%). Such strong transcriptional asymmetry in the preference of G+T over A+C is typical for other plants. No difference for G+T versus A+C composition has been detected for fruit fly and worm introns. A complex mosaic of MRI patterns and their fundamental importance Intricate arrangement of genomic MRI patterns Different MRI regions are not randomly arranged relative to each other  ADDIN EN.CITE Bechtel200850750750732Bechtel, Jason MFedorov ACharacterization of Genomic Mid-Range InhomogenityHealth Science Campus97MS2008ToledoUniversity of Toledo23(Bechtel 2008). For example, Figure 6 illustrates that G+C-rich regions tend to be associated in clusters. On the other hand, the distribution of A+T-rich regions is much more close to a random distribution with the exception that A+T-rich regions avoid very close proximity to each other  ADDIN EN.CITE Bechtel200850750750732Bechtel, Jason MFedorov ACharacterization of Genomic Mid-Range InhomogenityHealth Science Campus97MS2008ToledoUniversity of Toledo23(Bechtel 2008). So far, investigators have examined only individual genomic patterns. The mutual arrangement of various genomic mid-range patterns has never been thoroughly investigated yet. Our preliminary results suggest that within mammalian genomes there is a complex mosaic picture of MRI regions. Modeling sequences only with one particular type of MRI compositional bias using MRI-generator program from the Genomic MRI package has proven not to be a trivial computational task  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Bechtel et al. 2008). This has given us an appreciation that the reconstruction of the entire set of MRI patterns in modeling DNA sequences is an extremely challenging mission due to a complex multi-layer non-randomness in genomic sequences. In addition, genomic sequences have an intricate organization of nested patterns and also with respect to the clustering of particular patterns. Some features of this complex organization were described as genomic fractals in several publications  ADDIN EN.CITE  ADDIN EN.CITE.DATA (Havlin et al. 1995; Cheng, Tong, and Zhang 2007; Pellionisz 2008). This arrangement has been studied by methods such as "detrended fluctuation analysis" and a "Brownian walk" to uncover relationships such as power law correlations and exponential decays, which assess the scaling behavior of a system. This scaling behavior is related to fractal geometry and deals with "self-similarity," defined as the property of resembling a subset of oneself. Earlier investigations of this kind generally confined themselves to clusters of purines and pyrimidines, but later studies have shifted to examining G+C and A+T clusters for the thermodynamic implications of their pair-binding  ADDIN EN.CITE Peng1992303017Peng, C. K.Buldyrev, S. V.Goldberger, A. L.Havlin, S.Sciortino, F.Simons, M.Stanley, H. E.Center for Polymer Studies, Boston University, Massachusetts 02215.Long-range correlations in nucleotide sequencesNatureNature168-703566365AnimalsBase SequenceDNA/ chemistryHumansIntronsNucleotide MappingRegulatory Sequences, Nucleic AcidStatistics as TopicNASA Discipline CardiopulmonaryNon-NASA Center1992Mar 120028-0836 (Print)1301010engPeng1995292917Peng, C. K.Buldyrev, S. V.Goldberger, A. L.Havlin, S.Mantegna, R. N.Simons, M.Stanley, H. E.Cardiovascular Division, Harvard Medical School, Boston, MA 02215, USA.Statistical properties of DNA sequencesPhysica A180-92221AlgorithmsAnimalsBase SequenceDNA/ chemistryData Interpretation, StatisticalEscherichia coliFractalsHumansIntronsInvertebratesLinguisticsNucleotide MappingSequence Analysis, DNA/ statistics & numerical dataSoftwareYeastsNASA Discipline CardiopulmonaryNon-NASA Center19950378-4371 (Print)11540495engHavlin1995272717Havlin, S.Buldyrev, S. V.Goldberger, A. L.Mantegna, R. N.Peng, C. K.Simons, M.Stanley, H. E.Center for Polymer Studies and Department of Physics, Boston University, MA 02215, USA.Statistical and linguistic features of DNA sequencesFractals269-8432AlgorithmsAnimalsBase SequenceChromosome WalkingChromosomes, FungalDnaDNA, FungalDNA, PlantData Interpretation, StatisticalEntropyExonsFractalsHumansIntronsInvertebratesLinguisticsModels, GeneticPurine NucleotidesPyrimidine NucleotidesSequence Analysis, DNA/ statistics & numerical dataNASA Discipline CardiopulmonaryNASA Discipline Number 14-10NASA Program Space Physiology and CountermeasuresNon-NASA Center1995Jun0218-348X (Print)11539281engHaring2001323217Haring, D.Kypr, J.Institute of Biophysics, Academy of Sciences of the Czech Republic, Brno.Mosaic structure of the DNA molecules of the human chromosomes 21 and 22Mol Biol Rep9-17281Chromosomes, Human, Pair 21Chromosomes, Human, Pair 22DNA/ chemistryDNA, Bacterial/chemistryEscherichia coli/geneticsHumansNucleic Acid ConformationRepetitive Sequences, Nucleic Acid2001Mar0301-4851 (Print)11710567engNicolay2004313117Nicolay, S.Argoul, F.Touchon, M.d'Aubenton-Carafa, Y.Thermes, C.Arneodo, A.Laboratoire de Physique, Ecole Normale Superieure de Lyon, 46 Allee d'Italie, 69364 Lyon Cedex 07, France.Low frequency rhythms in human DNA sequences: a key to the organization of gene location and orientation?Phys Rev Lett1081019310AT Rich SequenceBase Composition/geneticsBase SequenceBiological ClocksChromosome Mapping/ methodsComputer SimulationDNA/ chemistry/ geneticsGenome, HumanHumansModels, GeneticMolecular Sequence DataNonlinear DynamicsPeriodicitySequence Analysis, DNA/ methods2004Sep 30031-9007 (Print)15447453engCheng2005252517Cheng, J.Zhang, L. X.Department of Physics, Jinhua University, Jinhua 321017, China. Jh_Chengjun@163.comStatistical properties of nucleotide clusters in DNA sequencesJ Zhejiang Univ Sci B408-1265AnimalsBase CompositionBase SequenceChromosomes/geneticsDNA, Protozoan/ geneticsGenome, ProtozoanGenomicsNucleotides/ geneticsPlasmodium falciparum/ genetics2005May1673-1581 (Print)15822156engCheng2007262617Cheng, J.Tong, Z. S.Zhang, L. X.Department of Physics, Jinhua College of Profession and Technology, Jinhua 321017, China. jh_chengjun@163.comScaling behavior of nucleotide cluster in DNA sequencesJ Zhejiang Univ Sci B359-6485Base SequenceComputer SimulationDNA, Fungal/ geneticsEncephalitozoon cuniculi/ geneticsEvolution, MolecularModels, GeneticMolecular Sequence DataMultigene Family/ geneticsNucleotides/ geneticsSequence Analysis, DNA/ methods2007May1673-1581 (Print)17542065eng(Peng et al. 1992; Havlin et al. 1995; Peng et al. 1995; Haring and Kypr 2001; Nicolay et al. 2004; Cheng and Zhang 2005; Cheng, Tong, and Zhang 2007). The purpose of MRI regions Often, in the popular literature, genomes are presented as a set of texts or instructions. Such a representation implies that there should be an intelligent creature somewhere inside a cell interpreting these DNA texts. Thus, it is more appropriate to compare genomes with self-realization programs that autonomously fulfill their tasks and are able to respond on environment signals and conditions. Such programs must be extremely complicated for complex organisms, like humans, which are built from trillions of cells of hundreds of different kinds, yet sharing the same genomic sequence. There must be fundamental principles for construction and functioning of genomic programs. One of the most important is the Principle of Recursive Genome Function (PRGF) illuminated by Pellionisz  ADDIN EN.CITE Pellionisz200847947947917Pellionisz, A. J.HelixoMetry, HQ, 935 Rosette Court, Sunnyvale, CA 94086, USA. pellionisz@junkdna.comThe principle of recursive genome functionCerebellumCerebellum348-59732008/06/21AlgorithmsCerebellum/*physiologyDNA/chemistry/*geneticsGenes/*physiologyGenetic Code*Genome, Human*Genomics/methods/trendsHumansModels, NeurologicalNerve Tissue Proteins/geneticsPurkinje Cells/physiologyRNA/genetics20081473-4230 (Electronic)18566877http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1856687710.1007/s12311-008-0035-yeng(Pellionisz 2008). The author considers the genome as an unsupervised operating system. The well-known examples of such a system are neural networks for which mathematical models describing their behavior has been developed. According to Pellionisz, the recursive genome function is a process when at every step of development already-built proteins iteratively access sets of primary and ensuing auxiliary information packets of DNA to build constantly developing hierarchies of protein structures. In other words, there is a crucial flow of information from proteins back to the genomic DNA. According to Pellionisz, this principle converts a genome from a closed to an open physical system and resolves the paradox of genomic entropy posed by John Sanford  ADDIN EN.CITE Sanford20055055055056John C. SanfordGenetic Entropy & the Mystery of the Genome20051599190028(Sanford 2005). This perspective elucidates the importance of MRI regions as specific sites for changing genomic information by proteins. Indeed, MRI regions are intricately associated with unusual DNA conformations, which in turn are binding sites for a number of proteins. These proteins could stabilize and/or initiate DNA conformation transformation and propagate the signal along neighboring DNA segments. For instance, Z-DNA binding proteins could initiate this transformation from right-handed B-DNA to the left-handed Z-form. This structural transition changes the DNA supercoiling for the regional DNA landscape and additionally creates specific B-Z-boundaries with flipped-over bases. Such transformation could modify, open, and/or hide, some information on the genomic DNA not only at the protein binding site but within neighboring regions. Conclusions Overall, within vast areas of previously thought junk DNA, represented by introns and intergenic sequences, there exist an intricate mosaic of various MRI regions with extreme base compositions. Various genomic MRI regions are tightly associated with unusual DNA conformations and must be of crucial importance for proper functioning of multi-cellular eukaryotes. Understanding of genomic MRI functions is critical for the newly emerged field of Personal Genomics and also for drug discovery. Acknowledgment This material is based upon work supported by the National Science Foundation under Grant No. 0643542. FIGURE LEGENDS  Figure 1. Cartoon of 3D-structure of anti-parallel DNA duplex 5`TGGAATGGAA:TGGAATGGAA3` formed by two copies of TGGAA pentamer repeat. This picture is a snapshot of the structure with the identifier 103D obtained from the Protein Data Bank. The structure was resolved via solution NMR approach by Chow and co-authors  ADDIN EN.CITE Chou199430303017Chou, S. H.Cheng, J. W.Fedoroff, O.Reid, B. R.Howard Hughes Medical Institute, University of Washington, Seattle 98195.DNA sequence GCGAATGAGC containing the human centromere core sequence GAAT forms a self-complementary duplex with sheared G.A pairs in solutionJ Mol BiolJ Mol Biol467-7924131994/08/19Base CompositionBase SequenceCentromere/*chemistryHumansMagnetic Resonance SpectroscopyModels, MolecularMolecular Sequence DataMolecular StructureNucleic Acid ConformationNucleic Acid Heteroduplexes/*chemistryOligodeoxyribonucleotides/chemical synthesis/*chemistry1994Aug 190022-2836 (Print)8064859http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8064859S0022-2836(84)71521-X [pii] 10.1006/jmbi.1994.1521eng(Chou et al. 1994). Four arrows point to unpaired guanosine residues stacked between Hoogsteen G-A pairs.  Figure 2. Cartoon of 3D-structure of a C-rich strand fragment of the human centromeric satellite III d(CCATTCCATTCCTTTCC) that forms intra-molecurlar i-motif structure with C.C(+) pairs from parallel strands intercalated head-to-tail. This picture is a snapshot of the structure with the identifier 1G22 obtained from the Protein Data Bank. The structure was resolved via solution NMR approach by  ADDIN EN.CITE Nonin-Lecomte200171717117Nonin-Lecomte, S.Leroy, J. L.PMC Groupe de Biophysique de l'Ecole Polytechnique et de l'UMR 7643 du CNRS, 91128, Palaiseau, France. sn@pmc.polytechnique.frStructure of a C-rich strand fragment of the human centromeric satellite III: a pH-dependent intercalation topologyJ Mol BiolJ Mol Biol491-50630922001/05/24Base SequenceCentromere/*geneticsCytosine/chemistry/*metabolismDNA, Satellite/*chemistry/genetics/*metabolismHumansHydrogen-Ion ConcentrationIntercalating Agents/chemistry/*metabolismModels, MolecularNuclear Magnetic Resonance, Biomolecular*Nucleic Acid ConformationOligonucleotides/chemistry/genetics/metabolismProtons2001Jun 10022-2836 (Print)11371167http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1137116710.1006/jmbi.2001.4679 S0022-2836(01)94679-0 [pii]eng(Nonin-Lecomte and Leroy 2001) for uridine derivative methylated on the first cytidine base, d(5mCCATTCCAUTCCUTTCC), whose proton spectrum is better resolved. Modified residue 5-METHYL-2'-DEOXYCYTIDINE is demonstrated with white, blue, red, and grey spheres.  Figure 3. Cartoon of 3D-structure of a parallel-stranded G-quadruplex DNA formed by the Tetrahymena telomeric sequence d(T-T-G-G-G-G-T). This picture is a snapshot of the structure with the identifier 139D obtained from the Protein Data Bank. The structure was resolved by combined NMR-computational approach by  ADDIN EN.CITE Wang199348248248217Wang, Y.Patel, D. J.Department of Biochemistry and Molecular Biophysics, College of Physicians and Surgeons, Columbia University, New York, NY 10032.Solution structure of a parallel-stranded G-quadruplex DNAJ Mol BiolJ Mol Biol1171-8323441993/12/20DNA/chemistry/*ultrastructureHydrogen BondingMagnetic Resonance SpectroscopyModels, Molecular*Nucleic Acid ConformationOligodeoxyribonucleotides/chemistrySolutions1993Dec 200022-2836 (Print)8263919http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8263919S0022-2836(83)71668-2 [pii] 10.1006/jmbi.1993.1668eng(Wang and Patel 1993). Four strands are shown in yellow, pink, green, and blue.  Figure 4 shows the graphical output of the MRI-analyzer program for the first intron of the Dystrophin gene (marked as intron) and also for the SRI-generated random sequence based on the tetramer oligonucleotide frequency table of the intron (marked as random). The entire sequence of the 319 kb intron and the random sequence is displayed on the x-axis. Blue bars represent positions content-rich MRI regions on the sequence. Red bars represent content-poor MRI regions. The y-axis contains upper and lower thresholds for the given content type. (A) Genomic MRI analysis of A+G-rich and A+G-poor (or T+C-rich) regions; (B) Genomic MRI analysis of G+T-rich and G+T-poor (or C+A-rich) regions.  Figure 5. Cartoon of 3D-structure of a purine.purine.pyrimidine DNA triplex containing G.GC and T.AT triples. This picture is a snapshot of the structure with the identifier 134D obtained from the Protein Data Bank. The structure was resolved using a combined NMR and molecular dynamics approach by  ADDIN EN.CITE Radhakrishnan199348848848817Radhakrishnan, I.Patel, D. J.Department of Biochemistry and Molecular Biophysics, College of Physicians and Surgeons, Columbia University, New York NY 10032.Solution structure of a purine.purine.pyrimidine DNA triplex containing G.GC and T.AT triplesStructureStructure135-52121993/10/15AdenineBase CompositionBase SequenceComputer GraphicsCytosineDNA/*chemistryGuanineHydrogen BondingMacromolecular SubstancesMagnetic Resonance Spectroscopy/methodsModels, MolecularMolecular Sequence Data*Nucleic Acid ConformationOligodeoxyribonucleotides/*chemistryPurinesPyrimidinesThymine1993Oct 150969-2126 (Print)8069626http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8069626eng(Radhakrishnan and Patel 1993).  Figure 6. Visualization of G+C-rich (blue) and A+T-rich (red) MRI features in human introns using a 400 nt base window size. The scale for each sequence is independent and is given in its sub-heading in nucleotides per pixel. 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Watson Research Center, P.O. Box 218, Yorktown Heights, NY 10598, USA. rigoutso@us.ibm.comShort blocks from the noncoding parts of the human genome have instances within nearly all known genes and relate to biological processesProc Natl Acad Sci U S AProc Natl Acad Sci U S A6605-10103172006/04/263' Untranslated Regions5' Untranslated RegionsAnimalsBase SequenceConserved SequenceDNA/genetics*Genome, HumanHumansIntronsMolecular Sequence DataOpen Reading FramesRNA Interference*Untranslated Regions2006Apr 250027-8424 (Print) 0027-8424 (Linking)16636294http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1663629414475210601688103 [pii] 10.1073/pnas.0601688103engTsirigos200852952952917Tsirigos, A.Rigoutsos, I.Bioinformatics and Pattern Discovery Group, IBM Thomas J. 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X.Department of Physics, Jinhua College of Profession and Technology, Jinhua 321017, China. jh_chengjun@163.comScaling behavior of nucleotide cluster in DNA sequencesJ Zhejiang Univ Sci B359-6485Base SequenceComputer SimulationDNA, Fungal/ geneticsEncephalitozoon cuniculi/ geneticsEvolution, MolecularModels, GeneticMolecular Sequence DataMultigene Family/ geneticsNucleotides/ geneticsSequence Analysis, DNA/ methods2007May1673-1581 (Print)17542065engPellionisz200847947947917Pellionisz, A. J.HelixoMetry, HQ, 935 Rosette Court, Sunnyvale, CA 94086, USA. pellionisz@junkdna.comThe principle of recursive genome functionCerebellumCerebellum348-59732008/06/21AlgorithmsCerebellum/*physiologyDNA/chemistry/*geneticsGenes/*physiologyGenetic Code*Genome, Human*Genomics/methods/trendsHumansModels, NeurologicalNerve Tissue Proteins/geneticsPurkinje Cells/physiologyRNA/genetics20081473-4230 (Electronic)18566877http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1856687710.1007/s12311-008-0035-yengXDd@80*  A"?*Fig1_MRIpatternsPicture 6Fig1_MRIpatterns.tifbJ_)@bI'm& n_)@bI'mPNG  IHDR_wsRGBgAMA a cHRMz&u0`:pQ<IDATx^Gl\YP@"7Ox@OzCQ}]ՕJL)啩L92N{2>"轑HHHHHHHk'0$@$@$@$@$@$@$>AΏHHHHHH `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$     =@$@$@$@$@$@$(]b9I       `$@$@$@$@$@$@.A%$ I`߿s8'/_1Տ:T_g:y8'g6OC$@$@P  bJ:^ ):< L5 & WG`gPX9f LBNl5%睊ex']R'!1>-+zp?A$@$@I#g$@$@cg;I;ݧl"Ww"|_` MᦌD$@$@0  &0(x:eY;_{%x_.=#wqL>yŰwJ$;Op}or$@$@SNxʑ$@$ERہl۱dSx$$/y-eϕhU x*W'~ ېz0 [bsݣg"_ ~ U$ *jx$@$0 ( qz]ދՏwc/-_M^o:a{6~~:>4zbs&u*6!u_9[%m+RkSxM}MhmѢb%հEUغ| I\WΊH>΃gqt,~+鼌ڮ+j<'  ӍGTbXs=?> a9PZiC  ׸ | ADk;_·et@MWWv>{QFB=6Cc0iZf[~}(ܦKGJt_?`xyj-!coһa؊FlxzbKf-=s=,^ZUdyY퍠 dDYo98ScVö38q|&Cǟ2$@$@$0 ]IH% ˠs0*fwѼ>Î< :^޿&ވuFDпa#Z%*Y`wPZr`nNS܋u(j.֣UDTU4"+ G9 {3/iSTM c5||C2g^%^1 6i <-59- ᱣ{% W"@J͹ p&QQ ;KtC@󾧨em웄 sC\ydQ+ן-4vCeaWBKW*u];"?c;F m}qyAx,\;xdxq|{s琟@ˆX KbeJ] 44n‰'_ZE҃5/;(nHHF#@{HH`Fh{;niN}n9ou2rR^(75O( ZI#U]%T/]#'4j8j/IHEt7nBG΋uGG}kvCcc|}P+uJSuDJE " "Jx^ GNAcQWFUqXDZ.mAeeT^,*`9$F.@L UI .̈́XS,ה/Bd!=)&MϫAF~-*jvs^(? 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