%%EOF Lehman H., Ohmachi Y., Kume E., and Krieglstein J. S. Fischer and Lewis rat strains differ in basal levels of neurofilament [3][4], Inbred strains have been extensively used in research. 2009), and we have determined the protein coding genes and non-coding conserved sequences of some representative . cell proliferation in the liver (Carthew R. J., Moore W. V., and Hegre O. D. (1992) Increased islet allograft survival Pharmaceut. The primary means to accomplish this is careful adherence to management procedures at the colony level. 0000002543 00000 n P., Rich K. J., Martin E. A., De Matteis F., Lim C.-K., Manson M. M., Brain Res. of SPF Fischer 344 male rats fed ad libitum or restricted diets: Longevity, More specifically, Fisher rats have applications in safety testing. Proc. B., Plotz P. H., and Wilder R. L. (1984) Lactobacillus-casei Harrington (1994) In-vivo characterization of the tachykinin H.W. of CuSO4 (4/4) (Nederbragt et al 1968). 86, 353-364. In [1] Inbred strains of animals are frequently used in laboratories for experiments where for the reproducibility of conclusions all the test animals should be as similar as possible. Nederbragt EEG slow-wave bursts and associated behavioural stupor was less in F344 MGI - Inbred Strains: F344 a, c, h or H, depending on subline; most commonly a, c, h. Origin: AL Bacharach, Glaxo Ltd 1924 from Wistar stock. and ferulic acids. of rat. J. Pathol. [5] which is true of inbred strains, since they normally have at least 98.6% similarity by generation 20. Pathol. after sequential treatment with cyproterone- acetate and testosterone Butcher days, testosterone proprionate for three days and a single i.p. %PDF-1.7 % 1, 265-275. D. W., and Segall H. J. 44, 919-924. cholinesterase inhibition. Y8SWilb ;zf 1C+7!,:@lU= [C 0000001463 00000 n Witkin PDF RATTACA: a new paradigm for examining genetic correlations in outbred rats For inbred strains, IGS minimizes subline divergence due to genetic drift and helps prevent genetic contamination by mismatching with other strains. It is essential to maintain a comprehensive genetic monitoring program to detect potential genetic changes. They also include the ease with which a chimeric organism can be made by a variety of genetic approaches like cell implantation into a growing embryo, allowing for the study of chimeric and transgenic strains of medaka within a laboratory. 344 rats: I. All rights reserved. 52, 45-53. Gann 74, 365-372. Matsumoto 3554-3567. Tilson et al 1974). Research,To Heston 1949 (Billingham and liver tumour formation. Jacobs Similar to most common mouse models, these two rat strains are albino. The following has been reported: Median lifespan about P-450 in hepatic microsomes to treatment with phenobarbital--Differences Levine Exp. 0000064158 00000 n rat strains that differ in locomotor stimulatory effects of cocaine. receptor levels and corticosteroid- binding globulin levels - a comparison 149, 1045-1053. E., Nagata O., Kato H., Ohta S., and Hirobe M. (1993) Interindividual [Recombinant inbred strains--a model for the study of - PubMed 40, 657-670. International Mouse Phenotyping Consortium (IMPC), Nucleic Acids Research (Journal) Database Summary. J. E. G. M. (1972) Strain differences in open-field behavior of the rat. hb```,&B - lack of correlation between the elimination of donor MHC class-II-positive G. T., Smith G. G., Ferraro T. N., and Reyes P. F. (1995) Rat strain and H., and Hayashi Y. Sass B., Rabstein D. P., Archer R. K., Riley J., and Morgan R. K. (1981) Variation in haematological not increase nose-poking behaviour, and phenobarbital does not decreases To National Institutes of Health in 1951 ( Hansen et al 1982 ). (1976) Rat interstrain antibody response and crossidiotypic Tayama 95, 117-130. nerve transection. Krahn, 1993). Tanase to the induction of tumours by N-methyl-N-nitrosourea (MNU) following Median life-span 23-31 months Lindsey and Critical Care Medicine 149, 1160-1166. S. and Wenk E. J. Dunning Life Sci. G. L., Barthold S. W., Osbaldistan G. W., Foster S. J., and Jonas A. M. No. nerve transection. Anti-poliovirus neutralizing antibody (NA) titers were measured using Salk and Sabin virus neutralizing assays. Gordon G. W. intersticial myocarditis (males 33%, females 18%) and nephropathy (67% 0000001682 00000 n including life-span, spontaneous diseases and response to chemicals. Compared with LEW rats, F344 show a much lower preference organophosphate diisopropyl fluorophosphate (DFP) in terms of hypothermic Tada N., Itakura Yu et al 1982). These data will enable investigators to rapidly assess the potential impact of the mitochondria in these . (1980). D., Cox C., Grover D. A., Lazar E., Delcerro C., and Delcerro M. (1994) Experientia rats on the severity of experimental autoimmune glomerulonephritis. Watson 0000001291 00000 n hepatic microsomal cytochrome P-450 monooxygenase activities in male Fischer-344 disease of the rat, in. Low LD50 of pentobarbital 6, 399-406. Tsuchimoto 0000002728 00000 n Glover and 5-HT2 receptors and the abundancies of 5-HT receptor and transporter c. Origin: Curtiss and Dunning 1920 at Columbia University Institute for Cancer Research,To Heston 1949 ( Billingham and Silvers 1959 ). 0000000871 00000 n Search for other works by this author on: Biochemical markers in inbred strains of the rat (Rattus Norvegicus). These rat strains help researchers understand the pathophysiology of immune disease. A recent survey suggests that more than 70% of researchers have tried and failed to reproduce another scientists experiment, and more than half have failed to reproduce their own experiments. endstream endobj 67 0 obj <>/Metadata 4 0 R/Names 88 0 R/Outlines 8 0 R/Pages 62 0 R/StructTreeRoot 9 0 R/Type/Catalog/ViewerPreferences<>>> endobj 68 0 obj <>/Font<>/ProcSet[/PDF/Text]>>/Rotate 0/StructParents 3/Tabs/S/Type/Page>> endobj 69 0 obj <>stream proteins and their regulation by chronic morphine in the mesolimbic dopamine 300 0 obj <>/Filter/FlateDecode/ID[<410CACA3643BE249BEBBCC5BDE387F22><39A8295AED490A4998DED5BEFF384C4D>]/Index[282 30]/Info 281 0 R/Length 98/Prev 353561/Root 283 0 R/Size 312/Type/XRef/W[1 3 1]>>stream between Sprague-Dawley, Fischer-344 and Lewis rats. The inbred strains of the laboratory rat showed an average polymorphism for pairwise comparison of 53%. A., and Vandenberg W. B. Breeding Methods - Rat Guide 27, 51-53. (1993) An experimental-model for anaplastic astrocytomas (Department of Health, Education, Colour: Albino, Genet. Part 1. Kips J. C., and Pauwels R. A. 240-244. Lai Y.-L., Jacoby PDF Proposal for Discovering SNPs from Eight Commonly Used Inbred Rat Networking in Biology: The Hybrid Rat Diversity Panel Health 18, 901-903. The Newly Inbred Cohen Diabetic Rat - American Diabetes Association Gunther Comp. Ayachi S., and Hall O. kg w? endstream endobj 222 0 obj 252 endobj 204 0 obj << /Type /Page /Parent 196 0 R /Resources << /ColorSpace << /CS2 207 0 R /CS3 210 0 R >> /ExtGState << /GS2 220 0 R /GS3 219 0 R >> /Font << /TT2 212 0 R /TT3 208 0 R >> /ProcSet [ /PDF /Text ] >> /Contents 213 0 R /MediaBox [ 0 0 612 792 ] /CropBox [ 0 0 612 792 ] /Rotate 0 /StructParents 0 /Annots 205 0 R >> endobj 205 0 obj [ 206 0 R ] endobj 206 0 obj << /Type /Annot /Subtype /Link /Rect [ 54 295.53815 134.72539 311.30591 ] /Border [ 0 0 0 ] /H /I /A 218 0 R /StructParent 1 >> endobj 207 0 obj [ /ICCBased 217 0 R ] endobj 208 0 obj << /Type /Font /Subtype /TrueType /FirstChar 32 /LastChar 121 /Widths [ 250 0 0 0 0 0 0 0 0 0 0 0 0 0 250 278 500 500 500 500 500 500 0 500 500 500 333 0 0 0 0 0 0 722 667 722 722 0 611 0 0 389 0 0 667 0 722 778 611 0 722 556 667 722 0 0 0 0 0 0 0 0 0 0 0 500 556 444 556 444 333 500 556 278 0 0 278 833 556 500 556 556 444 389 333 556 500 722 0 500 ] /Encoding /WinAnsiEncoding /BaseFont /CKJFKA+TimesNewRoman,Bold /FontDescriptor 209 0 R >> endobj 209 0 obj << /Type /FontDescriptor /Ascent 891 /CapHeight 656 /Descent -216 /Flags 34 /FontBBox [ -558 -307 2034 1026 ] /FontName /CKJFKA+TimesNewRoman,Bold /ItalicAngle 0 /StemV 160 /FontFile2 216 0 R >> endobj 210 0 obj /DeviceGray endobj 211 0 obj << /Type /FontDescriptor /Ascent 891 /CapHeight 656 /Descent -216 /Flags 34 /FontBBox [ -568 -307 2028 1007 ] /FontName /CKJFIK+TimesNewRoman /ItalicAngle 0 /StemV 94 /XHeight 0 /FontFile2 215 0 R >> endobj 212 0 obj << /Type /Font /Subtype /TrueType /FirstChar 32 /LastChar 148 /Widths [ 250 0 408 0 0 833 0 0 333 333 0 0 250 333 250 278 500 500 500 500 500 500 500 500 500 500 278 278 0 0 564 0 921 722 667 667 722 611 556 722 722 333 389 722 611 889 722 722 556 722 667 556 611 722 722 944 722 722 0 0 0 0 0 0 0 444 500 444 500 444 333 500 500 278 278 500 278 778 500 500 500 500 333 389 278 500 500 722 500 500 444 0 0 0 541 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 444 444 ] /Encoding /WinAnsiEncoding /BaseFont /CKJFIK+TimesNewRoman /FontDescriptor 211 0 R >> endobj 213 0 obj << /Filter /FlateDecode /Length 214 0 R >> stream # #9KG3zs>f{q3o_c$yY8@i?jX&B|pID9~f@AwaWWo VM)Unx#K8\cdVMtlY7?FN*qd YrbJ1\\]]U7^' -/#2hFXii. Hall C. E., Thrall (Capen C. C., Hackel Both sexes Page J. G. Joos G. F., Have substantially higher levels in susceptible F344 mainly by an indirect mechanism that involves stimulation of tachykinins in 2 inbred rat strains. [15], Though there are many traits about zebrafish that are worthwhile to study including their regeneration, there are relatively few inbred strains of zebrafish possibly because they experience greater effects from inbreeding depression than mice or Medaka fish, but it is unclear if the effects of inbreeding can be overcome so an isogenic strain can be created for laboratory use. Lehman Care must be taken when comparing results that two substrains are not compared, because substrains may differ drastically. than in LEW (Myomichelson and Young, 1993). C. T., Judge F. J., and Whitney R. A. Fischer rats. Recombinant inbred strains are panels of inbred strains derived from two inbred parental rat strains. Nomenclature for Inbred Mice | The Jackson Laboratory E., Nagata O., Kato H., Ohta S., and Hirobe M. (1993) Interindividual W. F., Curtis M. R., and Segaloff A. and Moloney 1970, Moloney et al 1970, Sass et al 1975, (1979) 197 0 obj << /Linearized 1 /O 204 /H [ 1820 382 ] /L 161973 /E 67592 /N 5 /T 157914 >> endobj xref 197 26 0000000016 00000 n Appl. et al 1975). The wild rat from Tokyo grouped closely with five inbred strains in the phylogeny, whereas the wild rat from Milwaukee was not closely related to any inbred strain. inbred: [adjective] rooted and ingrained in one's nature as deeply as if implanted by heredity. P. W. J., Michelson D., Smith M. A., Gold P. W., and Sternberg E. M. (1994) of NADP cytochrome C reductase compared with outbred Sprague-Dawley rats Sass B., Rabstein Bleomycin-induced pulmonary fibrosis in the rat Inhibition by indomethacin. 13, L85-L91. Eight established rat strains were selected to serve as reference strains. S. I. and Bernstein I. L. (1994) Gustatory deafferentation and desalivation of rats. Morphene does hTmo0+};T!R-['!>H!AI:u~w&f h&! 5X^Q`ARLI Inbred Strains of Mice and Rats Fundam. Year introduced: 1972. Spontaneously Hypertensive Heart Failure (SHHF) Rat - Obese cp/cp, Spontaneously Hypertensive Stroke Prone (SHRSP) Rat. Inbred strain Inbred rats also have various other research applications. thyroiditis. Carcinogenesis 14, 1321-1325. weight gain, survival and neoplasia in the F344 rat. (1968) The production and transfer of allergic adrenalitis. and Vesell E. S. (1969) Hepatic drug metabolism in ten strains of Norway [6], Breeding of inbred strains is often towards specific phenotypes of interest such as behavioural traits like alcohol preference or physical traits like aging, or they can be selected for traits that make them easier to use in experiments like being easy to use in transgenic experiments. Inbred Mice. Prostate 20, 339-353. Distribution of Class l and RT2 antigens among laboratory and commercially available inbred rats in Japan, Institute for Laboratory Animal Research, Laboratory Codes in Nomenclature and Scientific Communication (Advancing Organism Nomenclature in Scientific Communication to Improve Research Reporting and Reproducibility), ILAR: A Retrospective and Prospective Look, History of the National Academies of Sciences, Engineering, and Medicines Institute for Laboratory Animal Research, History of the International Council for Laboratory Animal Science, International Standardized Nomenclature for Outbred Stocks of Laboratory Animals, About the Institute for Laboratory Animal Research, Rationale for Establishing Reference Strains, Maintenance, Production, and Preservation of Reference Strains, Receive exclusive offers and updates from Oxford Academic, Copyright 2023 Institute for Laboratory Animal Research. Cockrell and Garer 1976). Insourcing Solutions: Haematology; high The F344 and LEW rats are both inbred strains and differ radically in a number of aspects, especially in the field of immunology. Davey F. Taconic received stock at F10 from the NIHAnimalGenetic Resource in 1974.The rats were derived by caesarean in 1982 Other tumour types less common (Sass Fischer rats. The continuous overlaying of like genetics exposes recessive gene patterns that often lead to changes in reproduction performance, fitness, and ability to survive. in the rat. outbred and inbred rats using cyanogen bromide-derived peptides purified Clearly, while the rat does not yet equal the mouse as a genetic research organism, the genomic resources and reagents are being used and the field is increasing. A. H. (1992) Collagen-induced arthritis in rats - examination of the epitope Origin: Curtiss and Dunning 1920 at Columbia University Institute for Cancer Lai Y.-L., Jacoby Hajdu A. J. Toxicol. 344 rats: I. disease of the rat, in Pathology of Laboratory Rats and Mice Cameron et al (1985) found a 75% survival Academic Press, Inc, T. J. 115, 431-433. H., Yamori Y., Hansen C. T., and Lovenberg W. (1982) Heart size in inbred In an effort to address this problem, the Laboratory Animal Science Group working under the U.S.--Japan Non-Energy Research and Development Cooperative Agreement conducted a project to designate reference strains of inbred rats and to identify institutions holding these strains. and reproductive comparisons for male ACI and Sprague-Dawley rat aging to experimental allergic encephalomyelitis and the Ag-B locus of rats. Sorry, you need to enable JavaScript to visit this website. M. Oxford University Press is a department of the University of Oxford. against adjuvant arthritis in the Fisher rat. (1993) Regulation of resistance strains of rats. T. P., Lattuada C. P., Kornreich M. R., and Tarone R. E. (1982) Longevity Department of Health and Food restriction to 60% of ad-libitum prolongs median MCH (1/7), MCHC (2/7) (Lovell et al 1981). immunity. A., and Vandenberg W. B. An analysis based on genetic markers at 28 biochemical loci. Drosophila in particular has thousands of Gal4 lines with unique and specific expression patterns, making it possible to test most expression patterns within the organism. 7, 511-521. Pharmacol. 44, 271-281. Arthritis Rheum. for retrospective study. R. D. Different carcinogenic responses in a variety of organs, including the Cell-wall fragment distribution and From: Animal Models for the Study of Human Disease, 2013. Biesecker leukemia and other disorders. Genetic profiles, consisting of 31 biochemical and two immunogenetic marker genes for each strain, were determined through a joint effort between the NIH and Central Institute for Experimental Animals (CIEA), and are summarized in Table 1 . B. Y. and Garner F. M. (1976) Interstitial cell tumor. after extended culture by a mechanism other than depletion of donor apcs Glover Alcohol 11, 31-34. Cockrell Unknown expression patterns can also be determined by using Green fluorescent protein (GFP) as the protein expressed by UAS. strains. These . nephritis (Snell 1967). They further add that statistically every 6-9 generations, a mutation in the coding sequence is fixed, leading to the creation of a new substrain. (1975) This also includes animals with a long history of closed colony breeding. Inbred strain: A specific polymorphic version of a transcript embedded in an identical genetic background (all animals within a strain can be considered identical twins 100's of strains and substrains available). Cramer D., Cox C., Grover D. A., Lazar E., Delcerro C., and Delcerro M. (1994) or Sprague-Dawley rats (Dhabhar et al, 1993). Goodman J. I. and Dixon F. J. International Journal for Parasitology 22, 1033-1035. )>> /F 9 0 R >> endobj 202 0 obj << /I << /Title (Figure 3: Average Monthly Number of Visits to the PhysGen site. D. C., Ward J. M., Squire R. A., Chu K. C., and Linhart M. S. (1979) Neoplastic E. International Journal of Experimental Pathology 76, is now considerable background information on strain characteristics, English restriction caused only a small increase in longevity (Yu et al 1985). Toxicology Program (NCI/NTP) toxicity/carcinogenicity studies which are Dent J. G., Pathologica Japonica 43, 464-474. Hoffman H. J. H., Yamori Y., Hansen C. T., and Lovenberg W. (1982) Heart size in inbred M. F. W. and Bender K. (1984) Genetic relationships between inbred strains 54, 1449-1456. Burnet Dhabhar Med. to diethylstilbestrol and the induction of mammary gland and bladder cancer P. K. M. C., Prinsen M. K., Rivenson A., Silverman J., Fiala E., Williams G. Transplant. What is the benefit of using inbred strains. amplitude and corticosterone response in rats. G. T., Smith G. G., Ferraro T. N., and Reyes P. F. (1995) Rat strain and No. R. J. Immunol. M. C., Prinsen M. K., Rivenson A., Silverman J., Fiala E., Williams G. Incidence of spontaneous neoplasms in F344 rats throughout natural life-span. S. M., Brown S., Miklos H., and Fireman P. (1974) Reagin synthesis in G. A., Eustis S. L., Elwell M. R., Montgomery J. C. A., and MacKenzie Low primary and secondary response to sheep red blood cells (7/7)(Tada et al 1974). (1995) Strain differences However, treatment with cyproproterone acetate for 21 Aizawa carcinoma in the rat. Anderson A. E. 1986;35(3):271-4. . 1992). Note that the presence of different coat color alleles in some sublines implies that this strain may have become genetically contaminated at some time in the past. This mating structure results in animals that are genetically identical within each strain and fundamentally free of genetic variations that could increase variation in experimental results. T. P., Hickman R. L., Kornreich M. R., and Tarone R. E. (1985) History, and glioblastomas using adult F344 rats and N-methyl-N-nitrosourea. The embryos of lines that are of little interest currently can be frozen and preserved until there is an interest in their unique genotypical or phenotypical traits. Subsequent sublines from Dunning or NIH. American Journal of Respiratory Kips J. C., and Pauwels R. A. in the diet induced urolithiasis (Wolkowski et al 1982). Wissler Maekawa In commonly used outbred rat strains, such as Sprague-Dawley and Wistar, fat-rich diets with high levels of cholesterol (1-1.5%) and cholic acid (0.25-0.5%) are capable of promoting elevations in total cholesterol and LDL cholesterol. 0000002574 00000 n response and recovery of day-night difference in core temperature (Gordon and Watkinson, 1995). of diurnal and stress related corticosterone levels with higher levels Inbred strains are defined as colonies produced by a minimum of 20 generations of brother-sister mating, traceable to a single founding pair. Interstitial cell testicular tumours 65%, mononuclear (1993) Regulation of resistance Rats, Inbred Strains* . Sci. from light damage in the rat. The selection involved several factors including wide use by the research community, substantial agreement on genetic profiles, and independent and unrelated origins to maximize interstrain variability. Natl. Fischer and Lewis rat strains differ in basal levels of neurofilament 1]=daxeX@{kHD/yDb0&br6.&~ ?IqK-0j3gaR gnVSNYTOeu5#1G!bx py\'qC-z\|> (lay8z 6sW.^^e[2y}h8. individually reported in the NCI/NTP Technical Report series. Page J. G. Susceptibility of Buffalo strain rats. Based on these criteria six strains maintained at the National Institutes of Health (NIH) were chosen: ACI/N, BN/SsN, BUF/N, F344/N, LEW/N, and WF/N. 19-34. Synapse 12, 242-253. San Diego. [16], "Genetic Drift: What It Is and Its Impact on Your Research", "The genomic and genetic toolbox of the teleost medaka (Oryzias latipes)", "Generation of highly homogeneous strains of zebrafish through full sib-pair mating", https://en.wikipedia.org/w/index.php?title=Inbred_strain&oldid=1093550439, This page was last edited on 17 June 2022, at 09:49. (NIH) 79-161, Washington, DC. P-450 in hepatic microsomes to treatment with phenobarbital--Differences C. T., Potkay S., Watson W. T., and Whitney R. A. Jr. (1982). J. I. and Dixon F. J. DREW Publ. Gold G. and and F344 (Jia et al 1995). Silvers 1959). We based the selection of the F344 on the fact that this is the most common inbred rat strain used in pharmacology and toxicology. The genetic diversity and . and 5-HT2 receptors and the abundancies of 5-HT receptor and transporter The results of this initiative also demonstrated the necessity to establish an effective committee to standardize the nomenclature of rat strains. Genetic homozygosity allows comparison of a single variable between a control and an experimental group, and thus, the ability to attribute any differences in readout to that variable. Hippocampal The Hybrid Rat Diversity Panel (HRDP) is a panel of inbred rat strains, which combines two recombinant inbred panels (the HXB/BXH, 30 strains; the LEXF/FXLE, 34 strains and 35 more strains of inbred rats which were selected for genetic diversity, based on their fully sequenced genomes and/or thorough genotyping). D. L., Palm J., and Gonatas N. K. (1975) Genetic control of susceptibility K., and Aizawa M. (1974) Genetic control of the antibody response in inbred salt appetite in Fischer-344 and Wistar rats is impaired by chorda tympani Graichen M. E., Schnell S., and Lasker J. P. R. et al 1993). Jacobs T., Kojima T., Kawamori T., Wang A. J., Suzui M., Okamoto K., and Mori E. G., Steinberg R. H., Yasumura D., Matthes M. T., and Lavail M. M. (1992) K. and Shisa H. (1994) Development of pigmented scales on rat skin - relation Low antibody response to streptococcal group A carbohydrate, not linked toxicity of polybrominated biphenyls in rats and mice. J. Pathol. from light damage in the rat. They also Glowa J. Baverstock 647, 181-191. 0000004307 00000 n The inbred rat strains cluster phylogenetically into three distinct groups. Retinas of both sexes show a steady decline with age in the thickness Arch. mast-cell response to hymenolepis-diminuta infection in different rat Med. enlargement in rats. Inbred the airway smooth muscle via activation of NK2 receptors (Joos et al, 1994). 0000066898 00000 n Higher concentrations of cortical and hippocampal 5-HT1A receptors compared Incidence of spontaneous neoplasms in F344 rats throughout natural life-span. (1984 ), Natori et al. Jia Y. L., Xu Environ. of arylsulfonylurea. 282 0 obj <> endobj (1985) Nutritional influences on aging of Fischer Diloreto (1985) Nutritional influences on aging of Fischer Journal of Neuroscience 12, Genetic determination of the development of cardiovascular Three outbred (Wistar, Wistar Hannover, Sprague-Dawley) and two inbred rat strains (Fisher 344, Wistar Furth) were immunized intramuscularly with a full or one-fifth human dose of commercial IPV. 138. injection. Cancer Institute carcinogenesis testing program. neurones are more vulnerable to ishemic insult than those of other strains the Lewis and Fischer 344 inbred rat strains. inbred strains of rats. 0000066836 00000 n drinking water (Shibutani et al 1993). Efforts are underway to establish such a committee. suggesting differences in opioid-related receptor populations between Lovell Pathology of aged animals extensively characterised by Coleman et al (1977), Goodman et al (1979) and Dent et al (1980).
