Studies of Growth Hormone Gene

Sequence Studies

Byrne C. R, et al.1987 isolated and sequenced the ovine growth hormone, together with about 1 kbp of DNA flanking each end of the gene. The structure of the gene was similar to that found for other growth hormone genes, particularly the bovine gene, and has a primary transcript of 1792 bp, with five exons, and with intron sizes of 264 bp, 231 bp, 227 bp and 273 bp.

Orian et al. 1988 Sequenced the sheep GH gene and the coding sequence predicted from that corresponded to the cDNA sequence for pituitary GH, and found that this gene presumably codes for the hormone expressed in the pituitary.

Guron C, et al. 1992 prepared cDNA from the mRNA isolated from sheep anterior pituitary glands. On cloning cDNA in E. coli, a clone coding full sequence of sheep pre-growth hormone was determined. The sequence for the sheep growth hormone (GH) was in agreement with the amino acid sequence of the protein determined previously except positions 99 and 146.

Yamono Y, et al. 1988 isolated the cDNA that encodes goat growth hormone (gGH), from a goat pituitary cDNA library. The cDNA was about 880 base pairs long and could code for a polypeptide of 217 amino acids, The amino acid sequence homology between gGH and the sequences of bovine GH, rat GH and human GH was 99, 83 and 66%, respectively.

Woychik R. P., et al. 1982 Cloned and sequenced the bovine growth hormone gene. The sequence was about 1800 bp with four introns. The comparison of untanslated regions of the bovine, human and rat growth hormone genes reveled many areas of highly conserved sequence showing divergent evolution.

Wallis OC & Wallis M. 1995 Cloned and characterized rabbit growth hormone encoding gene. The amino acid sequence of rabbit GH was similar to that of pig GH and other conserved mammalian GH, but differed markedly from the available sequences of ruminant and primate GH. No evidence of cluster of GH like genes were found as found in primates. This provides further support for the idea that, in mammals, GH shows a slow underlying rate of evolution.

 

Duplication of GH Gene in Caprine Ruminants (Sheep and Goat)

Valinsky et al. 1990 concluded on the basis of studies on a restriction fragment length polymorphism (RFLP), that there were two alleles at the GH gene locus in sheep and goats. In sheep, in one allele (Gh1), the GH gene was represented by a single copy (GH1 gene), while in the other (Gh2) the GH gene was duplicated (GH2-N (5¢) and GH2-Z (3¢) genes). Restriction maps of the sheep Gh1 and Gh2 loci indicated that the GH1, GH2-N and GH2-Z genes were all very similar.  The sequences flanking the 5¢ ends of the GH1 and GH2-N genes were similar, but differ from that flanking the GH2-Z gene. Individual animals were homozygous for Gh1 (i.e. possessed 2 GH-like genes) or Gh2 (4 GH-like genes) or heterozygous, with one copy of Gh1 and one of Gh2 (3 GH-like genes). The frequency of the Gh2 allele was greater than that of the Gh1 allele.

Gootwine E. et al. 1993 found PvuII restriction fragment length polymorphism at the growth hormone locus in sheep carrying the GH2 allele where the gene was duplicated. By restriction analysis and using the polymerase chain reaction they demonstrated that this RFLP was due to a mutation at the PvuII site located in the second intron of the 3' copy of the GH2 allele.

Gootwine et al. 1996 concluded that, in an animal homozygous for the Gh2 allele, only the GH2-N gene was expressed in the sheep pituitary.

Ofir et al. 1997 Sequenced each of the GH1, GH2-N and GH2-Z copies of the ovine GH genes of Awassi (a fat tailed breed) and Romney breeds of sheep and compared them with DNA sequences of pituitary and placental ovine GH expressed. (Gene Bank Accession numbers AF002110-AF2129) The results showed that the structure of the single GH gene copy of the ovine GH1 allele was highly conserved and the structure of the GH2-N copy of the GH2 allele was slightly divergent from the GH1 copy and was polymorphic, and that the GH2-Z gene copies were also polymorphic and accumulated more substitutions both in the coding and the non-coding regions then the GH2-N gene copy.

Lacroix et al. 1996 studied growth hormones expressed in ovine placenta and found two GH proteins, 22 and 28 kD. Sequences for three GH related cDNAs derived from sheep placenta were also worked out. One of thoes, coding for a protein identical to pituitary preGH, corresponded to the product of the GH1 or GH2-N gene. The other two were very similar and were coding for a protein differing from pituitary preGH at four amino acid residues, which appeared to correspond to the product of the GH2-Z gene.

Yamano et al. 1991 isolated two clones containing goat growth hormone (gGH) genes from goat genomic library using goat growth hormone cDNA as a probe. One clone CgGH contained gGH1 gene, and another clone, EgGH, contained gGH2 and gGH3 genes in tandem. DNA fragments containing gGH1 gene, and that containing gGH2 and gGH3 genes, were estimated to be allelic on the goat chromosome.

GH Gene Duplication in Animals Other Than Caprine Ruminants

Seeburg PH. 1982 sequenced three non-allelic genomic DNA fragments with each containing one member of growth hormone gene. These genes were coding for the known polypeptide hormones, growth hormone (hGH), chorionic somatomammotropin (hCS), and a yet unknown protein, which differed from hGH in 13 positions. Each gene was structured into five exons, the four introns occurring at identical positions, reflecting recent gene divergence.

Barsh GS et al. 1983 determined the structure of the human growth hormone gene cluster over a 78-kilo base region of DNA by the study of two overlapping cosmids. There were two growth hormone genes interspersed with three chorionic somatomammotropin genes, all in the same transcriptional orientation. Analysis of the sequences of the genes and identification of at least three different classes of duplication units interspersed throughout the five gene cluster suggests that the cluster evolved quite recently and that the mechanism of gene duplication involved homologous but unequal exchange between middle repetitive elements of the Alu family.

Chakravarti A. et al. 1984 detected six RFLPs in human growth hormone / human somatomammotropin gene cluster, patterns of polymorphism and linkage disequilibrium suggested independent origins of the human growth hormone gene cluster.

Hirt H. et al. 1987 obtained genomic clones containing the closely related genes for human growth hormone (hGH) and chorionic somatomammotropin (hCS), from genomic bacteriophage lambda and cosmid libraries. The hGH/hCS locus contained two GH genes and three CS genes spanning 48 kb of DNA. Data obtained from that experiment combined with the nucleotide sequences of all five GH and CS genes, indicated that the hGH/hCS gene locus had evolved by duplication mechanism.

Chen et al. 1989 sequenced the human chromosomal growth hormone locus contained on cloned DNA and spanning approximately 66,500 bp, and it was found that this locus evolved by a series of duplications and was found to contain five genes, which displayed 95% sequence identity. The expression of each gene was examined by screening pituitary and placental cDNA libraries by using gene-specific oligonucleotides. According to this analysis, the hGH-N gene was transcribed exclusively in the pituitary, whereas the other four genes (hCS-L, hCS-A, hGH-V, hCS-B) were expressed only in placental tissue.

Bauman G. 1999 identified several isoforms of growth hormone (GH) in humans. There were many reasons for this heterogeneity. At the genetic level, two genes encode GH: GH-N, expressed in the pituitary, and GH-V, expressed in the placenta. At the mRNA level, GH-N undergo alternative splicing into 20K and 22K isoforms. Post-translationally, 22K GH undergo modifications, such as acetylation at its amino terminus, deamidation, and oligomerization. The picture was complicated further in the circulation, where GH binds to two GH-binding proteins, each with different affinities for the GH isoforms.

Golos et al.1993 screened Rhesus monkey pituitary and placental cDNA libraries for hCS-hybridizing clones and found that there were at least five genes that code for GH-like proteins, one expressed in the pituitary and four in the placenta, although these may not be exactly equivalent to the human GH-like genes.

Ber et al. 1993 reported the sequences of two growth hormone encoding genes from tilapia fish (Tilapia nilotica). Sequencing data indicated that it was a consequence of a relatively recent duplication event. The two genes were highly homologous, having a similar intron (five)/exon (six) arrangement, and both encode an identical polypeptide.

 

Physiological Effects of GH Copy Number

Fleming et al.1997 investigated growth hormone (GH) gene expression in pituitaries of lambs from flocks selected for high (fat) or low (lean) back fat depth, which were also homozygous for a single GH gene allele, heterozygous or homozygous for duplication in the GH gene. It was conclude that the pituitary glands of lean sheep were bigger and have an increased GH content, but appeared to contain similar concentrations of GH mRNA and immunoreactive GH as the pituitaries of fat sheep. The presence of the GH gene duplication in sheep had little measurable effect on the expression and storage of GH in the pituitary.

Gootwine et al. 1997 studied the effect of growth hormone gene copy number (two three or four for different genotypes of GH gene) on growth parameters, plasma GH profile and response to GHRH. No significant effect of GH genotype on any growth parameter or body composition was found. But Significant differences were found between GH genotype and response to GHRH, this suggested that polymorphism in GH gene copy number may have its physiological implications for the functioning of the GH axis.

 

Polymorphism Studies by PCR

Balatskii VN and Pocherniaev KF. 1995 investigated Restriction fragment length polymorphism of growth hormone gene in Big White and Mirgorod pigs. A polymorphous BsuRI restriction site was found within an amplified gene-region near 5-end. It was localized within the first intron, 1.6 b.p. higher than exon 2 was. Homozygous as well as heterozygous animals according to variants of the gene were found.

Fu J. et al. 1995 studied MspI polymorphism in THY1 gene, encoding a cell surface glycoprotein predominantly expressed in brain and peripheral nerves. Two primers derived from the sequences flanking the polymorphic MspI site in intron 2 of the human THY1 gene were selected for RCP to amplify a 566 bp fragment that spans the MspI polymorphism. Polymorphism was detected by MspI digestion of the PCR product.

Polymerase Chain Reaction

 

Primer Designing

Rychlik W. (1995 a) explained the importance of careful selection of primers as most important factor affecting the quality of PCR.  In general, the longer the PCR product the more difficult it was to select efficient primers and set appropriate designing primers, and in general, the more DNA sequence information was available, the better the chance of finding an optimal primer pair. Efficient primers could be designed by avoiding the following flaws: primer-dimer formation, self-complementarity, too low Tm of the primers, and/or their incorrect internal stability profile.

Nei M and Li WH. 1979 developed a mathematical model for the evolutionary change of restriction sites in mitochondrial DNA. Formulas based on this model were presented for estimating the number of nucleotide substitutions between two populations or species. To express the degree of polymorphism in a population at the nucleotide level, a measure called "nucleotide diversity" was proposed.

Bains W.1994 examined the constraints of the design of oligonucleotide probes that were meant to hybridize to different unique sites in human genomic DNA. In 522 kb of human genomic DNA, 75% of 12-base and 89% of 22-base were unique, as opposed to 90% and 100% as expected of unstructured DNA, and this was not due solely to repetitive elements in the DNA. Hybridization in TMAC to reduce A+T content effects on melting temperature allows only 90% of unique targets to be hybridized under one set of conditions if a 2 degrees C difference between matched and mismatched sequences was required. This suggested that probe; hybridization applications would be harder than previously suggested.

Kwok S. 1990 investigated the effects of various primer-template mismatches on DNA amplification of an HIV-1 gag region by PCR. Single internal mismatches had no significant effect on PCR product yield while those at the 3'-terminal base had varied effects. A:G, G:A and C:C mismatches reduced overall PCR product yield about 100-fold, A:A mismatches about 20-fold. All other 3'-terminal mismatches were efficiently amplified. It was noted that mismatches of T with either G, C, or T had a minimal effect on PCR product yield. Double mismatches reduced PCR product yield dramatically, however presence of a mismatched T at the 3'-terminus, allowed significant amplification even when coupled with an adjacent mismatch. Furthermore, even two mismatched Ts at the 3'-terminus allowed efficient amplification.

 

Bioinformatics Involved in PCR

Thompson JD. et al. 1994 improved the sensitivity of the commonly used progressive multiple sequence alignment method for the alignment of divergent protein sequences. Firstly, individual weights were assigned to each sequence in a partial alignment. Secondly, amino acid substitution matrices were varied at different alignment stages according to the divergence of the sequences to be aligned. Thirdly, residue-specific gap penalties and locally reduced gap penalties in hydrophilic regions encourage new gaps in potential loop regions rather than regular secondary structure. Fourthly, positions in early alignments where gaps have been opened receive locally reduced gap penalties to encourage the opening up of new gaps at these positions. These modifications were incorporated into a new program, CLUSTAL W that was freely available.

Thompson JD. et al. 1997 developed CLUSTAL X, a new windows interface for the multiple sequence alignment program CLUSTAL W. The new system was easy to use, providing an integrated system for performing multiple sequence and profile alignments and analysing the results. A versatile sequence colouring scheme allowed the user to highlight conserved features in the alignment. Pull-down menus provided all the options required for traditional multiple sequence and profile alignment. CLUSTAL X worked well on many operating systems.

Altschul SF et al. 1997 described the BLAST programs as widely used tools for searching protein and DNA databases for sequence similarities. The authors described a variety of statistical refinements, which decreased execution time of the BLAST programs while enhancing their sensitivity to weak similarities. A new criterion for triggering the extension of word hits, combined with a new heuristic for generating gapped alignments, yields a gapped BLAST program that runs at approximately three times the speed of the original. In addition, a method was introduced for automatically combining statistically significant alignments produced by BLAST into a position-specific score matrix, and searching the database using that matrix.

 

Template DNA

Gustafson at el. 1993 showed that lengthy denaturation times of template DNA ranging from 1 to 7 min at pH 7.0-8.0, that were often employed prior to the start of a PCR reaction, resulted in marked degradation of the template. That can result in a significant reduction in the yield of PCR products larger than 500 bp, by up to 99%. This effect was demonstrated for both complex genomic template DNA, and also for a 2691-bp linear piece of template DNA using both a rapid hot-air thermocycler and a conventional block thermocycler. This decrease in product yield was likely due to the increased degradation of the template or target DNA as a result of pre-amplification denaturation (PAD). Therefore (PAD) was not recommend for amplifying larger pieces of DNA, irrespective of the starting pH of the template solution.

Vazquez and Steinberg 1999 showed that genomic DNA preparations derived from mammalian cells can often exhibit poor template activity in PCR, particularly when carried out on target sequences present at low copy number. Using genomic DNA bearing SV40 sequences integrated into host chromosomal DNA at low copy number as a target, they show that template efficiency can be dramatically enhanced after treatment of the genomic template with restriction enzymes for varying periods of time. Results indicated that optimal digestion time varied for each enzyme.

 

Taq Polymerase

Chien A et al. 1976 purified a stable DNA polymerase (EC 2.7.7.7) with a temperature optimum of 80ºC from the extreme thermophile, Thermus aquaticus. The enzyme was free from phosphomonoesterase, phosphodiesterase and single-stranded exonuclease activities. Maximal activity of the enzyme requireed all four dNTPs and activated calf thymus DNA. An absolute requirement for divalent cation cofactor was satisfied by Mg2+ or to a lesser extent by Mn2+. The pH optimum was 8.0 in tris-HCl buffer. The molecular weight of the enzyme was estimated approximately 63,000 to 68,000. The elevated temperature requirement, small size, and lack of nuclease activity distinguished this polymerase from the DNA polymerase of Escherichia coli.

Kaledin AS et al. 1980 isolated a 62 kD DNA polymerase from the thermophylic bacteria Thermus aquaticus YT-1. The enzyme does not contain contaminant exo and endonuclease activities and had a temperature optimum on the DNA templates at 70 degrees and that on RNA matrices at 50 degrees. The maximal activity of the enzyme required the presence of bivalent cations (Mg2+ or Mn2+) 0,1-0,2 M KCl or NaCl, all dNTPs and template in the incubation mixture. The enzyme was active when "activated" DNA, poly (dA)-poly (dT), poly (dA)-oligo (dT) 10 and poly (rA)-oligo (dT)10 were used as templates and was inactive on the native and denatured DNAs as well as on the native molecules of RNA and poly (rC)-oligo (dG) 12-180.

Saiki RK et al. 1988 used a thermostable DNA polymerase in an in vitro DNA amplification procedure, the polymerase chain reaction. The enzyme, isolated from Thermus aquaticus, greatly simplified the procedure and, by enabling the amplification reaction to be performed at higher temperatures, significantly improved the specificity, yield, sensitivity, and length of products that could be amplified. Single-copy genomic sequences were amplified by a factor of more than 10 million with very high specificity, and DNA segments up to 2000 base pairs were readily amplified. In addition, the method was used to amplify and detect a target DNA molecule present only once in a sample of 10(5) cells.

 

PCR Additives

Betaine

Rees W.A., et al. 1993 showed that the amino acid analogue betaine shares with small tetraalkylammonium ions the ability to reduce or even eliminate the base pair composition dependence of DNA thermal melting transitions. The "isostabilizing" concentration of betaine (at which AT and GC base pairs were equally stable) was approximately 5.2 M. Betaine exerted its isostabilizing effect without appreciably altering the conformation of double-stranded DNA from the B form. The presence of > 5 M betaine also did not greatly changed the behavior of DNA as a polyelectrolyte; this lack of effect on electrostatic interactions was expected because betaine exists as a zwitterion near neutral pH.

Henke W., et al, 1997 showed that Betaine improved the co-amplification of the two alternatively spliced variants of the prostate-specific membrane antigen mRNA as well as the amplification of the coding cDNA region of c-jun. It was suggested that betaine improved the amplification of these genes by reducing the formation of secondary structure caused by GC-rich regions and, therefore, may be generally applicable to improve the amplification of GC-rich DNA sequences.

Bovine Serum Albumin

Waleed Abu Al-Soud and Peter Rådström in 2000 investigated the effects of 16 amplification facilitators to enhance DNA amplification in the presence of blood, feces, or meat. The addition of 0.6% (wt/vol) bovine serum albumin to reaction mixtures containing Taq DNA polymerase reduced the inhibitory effect of blood and allowed DNA amplification in the presence of 2% instead of 0.2% (vol/vol) blood. Taq DNA polymerase was able to amplify DNA in the presence of 4% instead of 0.4% (vol/vol) feces and 4% instead of 0.2% (vol/vol) meat, The single-stranded DNA binding T4 gene 32 protein (gp32) had a relieving effect similar to that of BSA, they also found that addition of 11.7% (wt/vol) betaine allowed Taq DNA polymerase to amplify DNA in the presence of 2% (vol/vol) blood, while the addition of proteinase inhibitors allowed DNA amplification in the presence of 4% (vol/vol) feces. When various combinations of betaine, BSA, gp32, and Proteinase inhibitors were tested, no synergistic or additive effect was observed. The effects of facilitators on real-time DNA synthesis instead of conventional PCR were also studied.

Kreader C.A.1996 studied the benefits of adding bovine serum albumin (BSA) or T4 gene 32 protein (gp32) to PCR, with reaction mixtures containing substances that inhibit amplification. Whereas 10 to 1,000 fold more FeCl3, hemin, fulvic acids, humic acids, tannic acids, or extracts from feces, freshwater, or marine water were accommodated in PCR when either 400 ng of BSA per microl or 150 ng of gp32 per microl was included in the reactions, neither BSA nor gp32 relieved interference significantly when minimum inhibitory levels of bile salts, bilirubin, EDTA, NaCl, sodium dodecyl sulfate, or Triton X-100 were present. Use of BSA and gp32 together offered no more relief of inhibition than either alone at its optimal level, and neither protein had any noticeable effect on amplification in the absence of inhibitors.

Paabo S. et al. 1988 amplified pieces of mitochondrial DNA from a 7000 year old human by the polymerase chain reaction and sequenced them. Albumin and high concentrations of polymerase were required to overcome a factor in the brain extract that inhibited amplification. For this and other sources of ancient DNA, authors found an extreme inverse dependence of the amplification efficiency on the length of the sequence to be amplified. This property of ancient DNA distinguished it from modern DNA and thus provided a new criterion of authenticity for use in research on ancient DNA.

Sulfoxide Additives

Certain organic solvents, such as DMSO and betaine, had been reported to enhance PCR amplification, particularly for hard to amplify high GC templates. Chakrabarti R. and Schutt CE. 2002 studied two groups of compounds amides and sulfones and discovered several potent PCR enhancers, often outperforming DMSO. They identified tetramethylene sulfoxide as the most potent sulfur-oxygen compound in the enhancement of PCR amplification and as one of the most potent PCR enhancers currently known.

Choi JS et al. 1999 described the use of dimethyl sulfoxide (DMSO) for sequencing of high GC content templates. It was found that 5% (v/v) of DMSO in the reaction mixture recovered sequencing signal intensity with reduced frequency of ambiguous bases. DMSO has been empirically introduced to enhance the efficiency of PCR in GC-rich templates. However, the underlying mechanism of improved cycle sequencing by DMSO was unknown. Most of methyl nitrogen ring containing chemicals did not improve sequencing accuracy, whereas only glycerol mimicked the positive effect of DMSO by the same extent.

Baskaran N. et al. 1996 developed a PCR method for uniform amplification of a mixture of DNA templates differing in GC content, using the two enzyme approach (Klentaq1 and Pfu DNA polymerase) and a combination of DMSO and betaine. This method was applied to amplify the CGG repeat region from the fragile X region.

Ammonium Sulphate

Watanabe M. et al. 1996 established a simple and reproducible assay of polymerase chain reaction (PCR), to detect trinucleotide repeat expansion for Huntington's disease (HD) using a new DNA polymerase and buffer system. The system consists of an extremely heat stable DNA polymerase (Pfu), and a buffer supplemented with ammonium sulfate and dimethyl sulfoxide. Previous methods to amplify expanded alleles for HD have been very complex in PCR conditions, and of low reproducibility because of repetitive sequences around the primer sequences. With the present method, strong bands for the disease alleles were easily and reproducibly visible.

Olive DM et al. 1989 established a PCR assay for direct detection of human cytomegalovirus (HCMV). The efficiency of the amplification reaction was examined by using three different buffers and concentrations of dNTPs. The PCR assay was most efficient with a reaction mixture containing 17 mM ammonium sulfate, 67 mM Tris hydrochloride (pH 8.5), 7 mM MgCl2, 10 mM 2-mercaptoethanol, 170 micrograms of bovine serum albumin per ml, and each dNTP at a final concentration of 1.5 mM. The viral DNA was successfully amplified directly from 5 microliters of urine preheated at 93ºC for 30 min.

Miscellaneous Additives

Demeke and Adams (1992) surveyed of the inhibitory effects of various plant polysaccharides on PCR amplification of a 974-bp section of rbcL in Spinach and revealed that most of the polysaccharides tested were not inhibitory. In contrast, two of the acidic polysaccharides (dextran sulfate and gum ghatti) were inhibitory. The addition of 0.5% Tween 20 reversed the inhibitory effects of gum ghatti (polysaccharide: DNA ratio of 500:1). The inhibitory effect of dextran sulfate (50:1) could be reversed by the addition of Tween 20 (0.25 or 0.5%) DMSO (5%), or polyethylene glycol 400 (5%) but none of these three additives were effective at a 100:1 ratio of dextran sulf ate/DNA.

Varadaraj K, Skinner DM. 1994 describe conditions that improve the specificity of amplification of a GC rich (57% G + C) DNA by PCR. Under standard conditions a 368-bp segment of the satellite DNA of the Bermuda land crab was not amplified specifically. To establish optimal conditions two enzymes, AmpliTaq DNA polymerase and AmpliTaq DNA polymerase, Stoffel fragment (SF) along with  and a number of co-solvents were studied. Reagents that improved specificity of the amplified product were formamide, glycerol, DMSO, Tween-20 and NP-40; on the other hand, urea, ethanol and 1-methyl-2-pyrrolidone (NMP) inhibited amplification. Of the two enzymes, SF was more specific and efficient.

Kovárová M. and Dráber P (2000) compared the ability of several Tetramethylammonium (TMA) derivatives and some other reagents to increase the specificity of PCR and to improve the yield of amplification. A novel combination of the enhancer TMA and oxalate as anion was demonstrated to be a powerful enhancer of PCR. Addition of 2 mM TMA oxalate to the PCR mixture decreased the formation of non-specific DNA fragments and increased the yield of specific PCR products.

 

Primer Annealing and Melting

Ishii K. and Fukui M. (2001) reduced PCR bias derived from a primer mismatch, the effect of the annealing temperature on the product ratio was investigated by denaturing gradient gel electrophoresis analysis of PCR products from a mixture of perfect-match and one-mismatch templates. These templates were generated by PCR from Pediococcus acidilactici for one mismatch and Micrococcus luteus for the perfect match. PCRs showed that the bias was reduced at lower temperatures.

Breslauer KJ. et al. 1986 reported the complete thermodynamic library of all 10 Watson-Crick DNA nearest-neighbor interactions. This thermodynamic data was used to calculate temperature dependent behavior of any DNA duplex from its sequence. Authors illustrated a method of calculation by using the nearest-neighbor data to predict transition enthalpies and free energies for a series of DNA oligomers. These predicted values were in excellent agreement with the corresponding values determined experimentally. This agreement demonstrated that a DNA duplex structure thermodynamically can be considered to be the sum of its nearest-neighbor interactions.

Peuschel KE. 2000 worked with megaprimers of length 450-770 bp, optimal annealing of the megaprimer to the template-DNA was shown to occur at unexpectedly low temperatures under conditions suboptimal for amplification of the elongated megaprimer product by the short flanking primers, although very high annealing temperatures had been expected to give best results. The finding contradicted the rule that annealing temperatures should be just below the melting temperature of a given primer, and leaded to the conclusion that the curves of melting and annealing temperatures diverge rather than running in parallel.

Owczarzy R. et al. 1997 compared the 11 published sets of nearest neighbor sequence dependent thermodynamic parameters for DNA, and demonstrated use of the nearest-neighbor sets in predicting tm from the DNA sequence. The authors also assessed the ability of the nearest neighbor parameters to provide accurate predictions of experimental tm's of short duplex DNA oligomers.

Rychlik W et al. 1990 experimentally determined the optimal annealing temperature (TaOPT) values for several primer template pairs and develop a method for its calculation. At both sub and super optimal Ta values, non-specific products may be formed, and the yield of products was reduced. Optimizing the Ta was especially critical when long products were synthesized or when total genomic DNA was the substrate for PCR. The TaOPT was found to be a function of the melting temperatures of the less stable primer-template pair and of the product. The fact that experimental and calculated TaOPT values agree to within 0.7 degree C eliminateed the need for determining TaOPT experimentally.

 

Contamination in PCR

Ou CY et al. 1991 described UV irradiation as a simple and efficient way to minimize contamination or false positivity in routine PCR tests. They characterized the effect of UV irradiation on DNA template, primers, deoxynucleoside triphosphate and Taq polymerase. Reduction of the HIV DNA templates in polypropylene microcentrifuge tubes by more than 1000-fold can be achieved by UV irradiation. The sensitivity of the primers was sequence and concentration dependent. Taq polymerase was highly UV sensitive, whereas deoxynucleotide triphosphate was relatively UV resistant.

Prince AM and Andrus L. 1992 investigated the use of sodium hypochloride (Clorox), in comparison to concentrated HCl, for PCR sterilization. Ten percent Clorox was found to eliminate all ethidium bromide-stainable DNA and to prevent PCR amplification of a 600-bp DNA segment within one minute of template treatment. RNA was similarly destroyed. By contrast, even 2.0 N HCl did not destroy DNA detectable by PCR within five minutes.

 

PCR Optimization

Linz U et al. 1990 optimized PCR parameters using different templates, together with 6 sets of primers, the following optimal conditions were found: the DNA should be linearized, primer concentration 0.1-0.2 mumol/L, magnesium ion concentration <2 mmol/L. pH 8.5-9.0. 25 cycles were sufficient. For fragments greater than 103 bases the elongation time should be 5 min. The elongation temperature was not found critical and could vary between 50 and 70ºC. The hybridization temperature was used to control the specificity of the polymerase chain reaction and, finally, mismatches at the 3' end of the primer could totally inhibit the amplification.

Rychlik W. (1995 b) tested effects of the primer concentration, annealing temperature, salt and solvent concentrations on PCR yield. Priming was detectable when the 3' terminal portion of the partially mismatched primer formed a continuous duplex more stable than -11 kcal/mol with the target DNA. In the presence of low magnesium ion concentrations, priming was significantly reduced; glycerol (5%) and formamide (2.5%) had only a slight effect (Taq DNA polymerase). Oligonucleotides that were GC rich at their 3' ends exhibited high priming efficiency but were also prone to false priming, since the shorter fragments of their 3' ends were stable enough to serve as primers.

Harris S and Jones DB. 1997 assessed the parameters, which influence DNA amplification efficiency and specificity. As no single protocol was appropriate for all situations and optimizing PCR required evaluation of the denaturation, annealing and extension temperatures, the number of cycles performed, and the primer, magnesium chloride, dNTP, Taq DNA polymerase and DNA template concentrations. The important parameters for efficient, specific amplification were denaturation time and temperature, stringent annealing temperatures and magnesium chloride concentration. The importance of DNA concentration was found to depend upon the source from which the DNA was extracted.

 

Restriction Enzymes

 

Star Activity

Nasri M et al. 1985 worked out the activity of Pvu II and found that the substrate specificity of PvuII endonuclease was relaxed in the presence of dimethyl sulfoxide. The new recognition sequences cleaved in pBR322 DNA have been found to be CCGCTG, CATCTG, CAGATG, CAGGTG and CAGCGG.

Nasri M. and Thomas D. 1987 found the restriction endonuclease PvuII that cleaves the sequence CAGCTG, decreased its substrate specificity in the presence of organic solvents. Thirty three sites were identified on the nucleotide sequence of pBR322 DNA. The new recognition sequences were shown to be AAGCTG, GAGCTG, CNGCTG, CANCTG, CAGNTG, CAGCNG, CAGCTC and CAGCTT. PvuII recognized and cleaved degenerate sequences, which differ from the standard PvuII sequence CAGCTG at only one of the recognition site. The optimum incubation medium for PvuII activity was found to be: 10-50 mM Tris-HCl, pH 8.5, 12-15 mM MgCl2, 50 mM NaCl, 10% ethanol + 10% dimethylsulfoxide.

George J and Chirikjian JG. 1982 studied the effect of glycerol on the specificity of DNA cleavage by the restriction endonuclease BamHI. In addition to the canonical G decreases from G-A-T-C-C site, BamHI cut DNA at several other sites, number of BamHI.1 sites in simian virus 40 and pBR322 was determined to be 13 for each DNA. Those included G decreases from G-A-A-C-C, G decreases from G-C-T-C-C, G decreases from G-G-T-C-C, and G-A-A-T-C-C with the complementary strand sequence assignments of G-G-T-T-C-C, G-G-A-G-C-C, G-G-A-C-C-C, and G-G-A-T-T-C. The relaxation in specificity was related to hydrogen bond acceptor and donor sites in the recognition sequence.

Nasri M. and Thomas D .1986 studied that under the standard reaction conditions, the restriction endonuclease HindIII cleaves double-stranded DNA, within the recognition sequence A decreases AGCTT. In the presence of Dimethyl sulfoxide the substrate specificity of this enzyme was reduced and cleavages occurred at additional sites. They had determined the secondary sites in pBR322 DNA under relaxed conditions and found that it cleaveed: G decreases AGCTT, A decreases GGCTT, A decreases TGCTT, A decreases ATCTT, A decreases AGCCT, A decreases AGCAT, A decreases AGCGT, A decreases AGCTC, producing fragments with cohesive ends.