Chromosome Banding Techniques

Rhona R. Schreck1, Christine M. Distèche2

1 Cedars‐Sinai Medical Center, Los Angeles, California, 2 University of Washington School of Medicine, Seattle, Washington
Publication Name:  Current Protocols in Human Genetics
Unit Number:  Unit 4.2
DOI:  10.1002/0471142905.hg0402s00
Online Posting Date:  May, 2001
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Abstract

Chromosome banding techniques produce a series of consistent landmarks along the length of metaphase chromosomes that allow for both recognition of individual chromosomes within a genome and identification of specific segments of individual chromosomes. These landmarks facilitate assessment of chromosome normalcy, identification of sites of chromosome breaks and alterations, and location of specific genes. This unit covers these basic banding techniques (Q‐banding, G‐banding, and R‐banding), which produce virtually identical patterns of bands along the length of human chromosomes, although the bands and polymorphic regions highlighted may differ with each technique. These techniques highlight reproducible landmarks along the length of the chromosome and specialized staining techniques can be used to highlight particular regions of chromosomes, such as heterochromatic and repeated‐sequence segments. These specialized techniques, nucleolar organizer region (NOR) staining, centromeric heterochromatin staining (C‐banding), methylated satellite DNA staining (distamycin‐DAPI banding), and replication banding are also presented in this unit.

     
 
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Table of Contents

  • Strategic Planning
  • Basic Protocol 1: Quinacrine Banding (Q‐Banding)
  • Reagents and Solutions for Q‐Banding
  • Commentary for Q‐banding
  • Giemsa Banding (G‐Banding)
  • Basic Protocol 2: GTG Technique for G‐Banding
  • Alternate Protocol 1: G‐Banding with Wright Stain
  • Support Protocol 1: Aging Slides with Heat
  • Support Protocol 2: Aging of Slides with Hydrogen Peroxide
  • Reagents and Solutions for G‐Banding
  • Commentary for G‐Banding
  • Reverse Banding(R‐Banding)
  • Basic Protocol 3: RHG Technique for R‐Banding
  • Alternate Protocol 2: CA3/DA Technique for R‐Banding
  • Reagents and Solutions for R‐Banding
  • Commentary for R‐Banding
  • Replication Banding
  • Basic Protocol 4: B‐Pulse Replication Banding for Lymphocytes
  • Alternate Protocol 3: T‐Pulse Replication Banding for Lymphocytes
  • Alternate Protocol 4: B‐Pulse Replication Banding for Fibroblasts
  • Alternate Protocol 5: T‐Pulse Replication Banding for Fibroblasts
  • Support Protocol 3: Visualization of BrdU Replication Banding by Fluorescent Dye
  • Support Protocol 4: Visualization of BrdU Replication Banding by Light Treatment and Giemsa Staining
  • Support Protocol 5: Visualization of BrdU Replication Banding by Heat Treatment and Giemsa Staining
  • Reagents and Solutions for Replication Banding
  • Commentary for Replication Banding
  • Basic Protocol 5: Centromeric Heterochromatin Staining (C‐Banding)
  • Reagents and Solutions for C‐Banding
  • Commentary for C‐Banding
  • Basic Protocol 6: Nucleolar‐Organizer‐Region Staining (NOR Staining)
  • Reagents and Solutions for NOR Staining
  • Commentary for NOR Staining
  • Distamycin‐DAPI Staining
  • Basic Protocol 7: Distamycin‐DAPI Staining
  • Alternate Protocol 6: Hoechst 33258–Distamycin Staining
  • Reagents and Solutions for Distamycin/DAPI Staining
  • Commentary for Distamycin/DAPI Staining
  • Sequential Staining of Slides
  • Basic Protocol 8: Sequential Staining
  • Support Protocol 6: Destaining
  • Commentary for Destaining
  • Figures
  • Tables
     
 
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Materials

Basic Protocol 1: Quinacrine Banding (Q‐Banding)

  MaterialsFor common stock solutions, see appendix 2D; for suppliers, see suppliers appendix.
  • Air‐dried slides of metaphase chromosomes (unit 4.1)
  • recipeQuinacrine staining solution (see recipe)
  • recipeMcIlvaine buffer, pH 5.6 (see recipe)
  • Immersion oil, low fluorescence
  • Coverslips, no. 0 or no. 1
CAUTION: Quinacrine is hazardous; see appendix 2A for guidelines on handling, storage, and disposal.

Basic Protocol 2: GTG Technique for G‐Banding

  MaterialsFor common stock solutions, see appendix 2D; for suppliers, see suppliers appendix.
  • HBSS ( appendix 2D)
  • recipeTrypsin solution (see recipe)
  • 70% and 90% (v/v) ethanol
  • recipe2% Giemsa staining solution (see recipe; prepare fresh)
  • Aged slides of metaphase chromosomes (see support protocols)
  • Xylene or Hemo‐De
CAUTION: Xylene is hazardous; see appendix 2A for guidelines on handling, storage, and disposal.NOTE: Because the stain is difficult to remove from skin, it is advisable to wear gloves when working with Giemsa.

Alternate Protocol 1: G‐Banding with Wright Stain

  Additional MaterialsFor common stock solutions, see appendix 2D; for suppliers, see suppliers appendix.
  • recipeBacto Trypsin solution (see recipe)
  • recipeSorensen phosphate buffer, pH 6.8 (see recipe)
  • recipeWright stain (see recipe; prepare fresh)

Support Protocol 1: Aging Slides with Heat

  Materials
  • Air‐dried slides of metaphase chromosomes (prepare freshly; unit 4.1)
  • 15% (v/v) H 2O 2 (dilute 30% H 2O 2 1:1 with water immediately before use)
  • 50°C hot plate or slide warmer
CAUTION: H 2O 2 is hazardous; see appendix 2A for guidelines on handling, storage, and disposal.

Support Protocol 2: Aging of Slides with Hydrogen Peroxide

  MaterialsFor common stock solutions, see appendix 2D; for suppliers, see suppliers appendix.
  • recipe1.25 M sodium phosphate buffer, pH 4.0 (see recipe)
  • Air‐dried slides of metaphase chromosomes (prepare fresh; unit 4.1)
  • recipe10% Giemsa staining solution (see recipe)
  • Coplin jars
  • 88°C circulating water bath with cover

Basic Protocol 3: RHG Technique for R‐Banding

  Additional MaterialsFor common stock solutions, see appendix 2D; for suppliers, see suppliers appendix.
  • recipeChromomycin A3 staining solution (see recipe)
  • recipeDistamycin A staining solution with MgCl 2 (see recipe)
  • Glycerol
  • Coverslips, no. 1
  • Additional reagents and equipment for fluorescence microscopy (unit 4.4)

Alternate Protocol 2: CA3/DA Technique for R‐Banding

  MaterialsFor common stock solutions, see appendix 2D; for suppliers, see suppliers appendix.
  • Phytohemagglutinin (PHA)‐treated lymphocyte cultures (unit 4.2)
  • 10 mM BrdU (30.7 mg in 10 ml H 2O; store in 1‐ml aliquots ≤6 months at −20°C)
  • 10 mM 2′‐deoxycytidine (dC; 28 mg in 10 ml H 2O; store in 1‐ml aliquots ≤6 months at −20°C)
  • 10 µg/ml Colcemid (e.g., GIBCO/BRL; store at 4°C)
  • Tissue culture vessel of desired size
  • Additional reagents and equipment for culture and metaphase harvest of peripheral blood (unit 4.1) and chromosome slide preparation (unit 4.1)
CAUTION: Human lymphocytes and BrdU are hazardous; see appendix 2A for guidelines on handling, storage, and disposal.NOTE: All incubations are performed in a humidified 37°C, 5% CO 2 incubator unless otherwise specified.

Basic Protocol 4: B‐Pulse Replication Banding for Lymphocytes

  Additional MaterialsFor common stock solutions, see appendix 2D; for suppliers, see suppliers appendix.
  • Tissue culture medium with and without 10 µM thymidine (from 1 mM thymidine stock; unit 8.3), 37°C
  • IEC HS‐NII centrifuge with 958 rotor (or equivalent)
NOTE: All incubations are performed in a humidified 37°C, 5% CO 2 incubator unless otherwise specified.

Alternate Protocol 3: T‐Pulse Replication Banding for Lymphocytes

  Additional MaterialsFor common stock solutions, see appendix 2D; for suppliers, see suppliers appendix.
  • Actively growing fibroblast cultures (∼8 to 10 hr after passage)
  • Tissue culture medium with and without 10 µM thymidine (from 1 mM thymidine stock; unit 8.3), 37°C
CAUTION: Human fibroblasts are hazardous; see appendix 2A for guidelines on handling, storage, and disposal.

Alternate Protocol 4: B‐Pulse Replication Banding for Fibroblasts

  MaterialsFor common stock solutions, see appendix 2D; for suppliers, see suppliers appendix.
  • Metaphase slides of BrdU‐substituted chromosomes (preceding basic or alternate protocols)
  • 95%, 70%, and 30% (v/v) ethanol
  • PBS ( appendix 2D)
  • recipeHoechst 33258 staining solution A (see recipe)
  • recipeMcIlvaine buffer, pH 7.5 (see recipe)
CAUTION: Hoechst 33258 is hazardous; see appendix 2A for guidelines on handling, storage, and disposal.

Alternate Protocol 5: T‐Pulse Replication Banding for Fibroblasts

  MaterialsFor common stock solutions, see appendix 2D; for suppliers, see suppliers appendix.
  • Metaphase slides of BrdU‐substituted chromosomes (preceding basic or alternate protocols)
  • PBS ( appendix 2D)
  • recipeHoechst 33258 staining solutions A and B (see recipe)
  • 2× SSC ( appendix 2D), 60° or 65°C
  • recipe4% Giemsa staining solution (see recipe)
  • 100 × 100 × 15–mm square plastic dishes with covers
  • 24 × 60–mm coverslips
  • 20‐W fluorescent light
  • 60° to 65°C water bath
CAUTION: Hoechst 33258 is hazardous; see appendix 2A for guidelines on handling, storage, and disposal.

Support Protocol 3: Visualization of BrdU Replication Banding by Fluorescent Dye

  MaterialsFor common stock solutions, see appendix 2D; for suppliers, see suppliers appendix.
  • Metaphase slides of BrdU‐substituted chromosomes (preceding basic or alternate protocols)
  • recipe1.0 M sodium phosphate buffer, pH 8.0 (see recipe)
  • recipe4% Giemsa staining solution (see recipe)
  • 87° to 90°C water bath

Support Protocol 4: Visualization of BrdU Replication Banding by Light Treatment and Giemsa Staining

  MaterialsFor common stock solutions, see appendix 2D; for suppliers, see suppliers appendix.
  • Air‐dried slides of metaphase chromosomes aged 1 week at room temperature (unit 4.1)
  • 0.2 M HCl
  • recipe5% (w/v) barium hydroxide [Ba(OH) 2; see recipe], 50°C
  • 2× SSC ( appendix 2D), 60°C
  • recipe4% Giemsa staining solution in Gurrs buffer (see recipe)
  • Gurrs buffer, pH 6.8: dissolve Gurrs pH 6.8 buffer tablets (Bio/medical Specialties) in water according to manufacturer's instructions
  • Glass and polyethylene Coplin jars
  • 50° and 60°C water baths

Support Protocol 5: Visualization of BrdU Replication Banding by Heat Treatment and Giemsa Staining

  MaterialsFor common stock solutions, see appendix 2D; for suppliers, see suppliers appendix.
  • Air‐dried slides of metaphase chromosomes (<7 days old, unheated; unit 4.1)
  • recipe2% gelatin solution (see recipe)
  • recipe50% silver nitrate solution (see recipe)
  • 3% (v/v) acetic acid (3 ml in 97 ml H 2O; prepare fresh)
  • 65°C slide warmer or drying oven
NOTE: Silver nitrate is a protein stain and will stain skin and clothing. Wear gloves and lab coat.

Basic Protocol 5: Centromeric Heterochromatin Staining (C‐Banding)

  MaterialsFor common stock solutions, see appendix 2D; for suppliers, see suppliers appendix.
  • Air‐dried slides of metaphase chromosomes (unit 4.1)
  • recipeMcIlvaine buffer, pH 7.0 and 7.5 (see recipe)
  • recipeDistamycin A staining solution (see recipe)
  • recipeDAPI staining solution (see recipe)
  • Immersion oil, low fluorescence
  • Humidified chamber (e.g., petri dish with moist paper towel)
  • Coverslips, no. 0 or no. 1

Basic Protocol 6: Nucleolar‐Organizer‐Region Staining (NOR Staining)

  Additional MaterialsFor common stock solutions, see appendix 2D; for suppliers, see suppliers appendix.
  • PBS ( appendix 2D)
  • recipeHoechst 33258 staining solution (see recipe)
  • Mounting buffer: 1:1 (v/v) glycerol/ recipeMcIlvaine buffer, pH 7.5

Basic Protocol 7: Distamycin‐DAPI Staining

  MaterialsFor common stock solutions, see appendix 2D; for suppliers, see suppliers appendix.
  • Stained slides
  • Xylene or Hemo‐De (Fisher)
  • 1:1 (v/v) xylene/ethanol (if xylene rather than Hemo‐De is used)
  • 95% ethanol
  • Fixative: 3:1 (v/v) methanol/acetic acid
  • Methanol (reagent grade)
CAUTION: Xylene is hazardous; see appendix 2A for guidelines on handling, storage, and disposal.
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Figures

Videos

Literature Cited

Literature Cited for Q‐Banding
   Caspersson, T., Farber, S., Foley, G.F., Kudynowski, J., Modest, E.J., Simonsson, E., Wagh, U., and Zech, L. 1968. Chemical differentiation along metaphase chromosomes. Exp. Cell Res. 49:219‐222.
   Caspersson, T., Zech, L., and Johansson, C. 1970. Analysis of human metaphase chromosomes by aid of DNA binding fluorescent agents. Exp. Cell Res. 62:490‐492.
   Dev, V.G., Warburton, D., Miller, O.J., Miller, D.A., Erlanger, B.F., and Beiser, S.M. 1972. Consistent pattern of banding of anti‐adenosine antibodies to human metaphase chromosomes. Exp. Cell Res. 74:288‐293.
   Distèche, C. and Bontemps, J. 1974. Chromosome regions containing DNAs of known base composition specifically evidenced by 2,7‐di‐t‐butyl proflavin. Chromosoma 47:263‐281.
   Distèche, C., Bontemps, J., Houssien, C., Frédérec, J., and Fredericq, E. 1980. Quantitative analysis of fluorescence profiles of chromosomes. Exp. Cell Res. 125:251‐264.
   Korenberg, J.R. and Rykowski, M.C. 1988. Human genome organization: Alu, Lines, and the molecular structure of metaphase chromosome bands. Cell 53:391‐400.
   Manuelidis, L. and Ward, D.C. 1984. Chromosomal and nuclear distribution of the Hin dIII 1.9 kb human DNA repeat segment. Chromosoma 91:28‐38.
   Pachmann, U. and Rigler, R. 1972. Quantum yield of acridines interacting with DNA of defined base sequence. A basis for the explanation of acridine bands in chromosomes. Exp. Cell Res. 72:602‐608.
   Schreck, R.R., Warburton, D., Miller, O.J., Beiser, S.M., and Erlanger, B.F. 1973. Chromosome structure as revealed by a combined chemical and immunochemical procedure. Proc. Natl. Acad. Sci. U.S.A. 70:804‐807.
   Schreck, R.R., Erlanger, B.F., and Miller, O.J. 1974. The use of anti‐nucleoside antibodies to probe the organization of chromosomes denatured by ultraviolet irradiation. Exp. Cell Res. 88:31‐39.
   Weisblum, B. and deHaseth, P.L. 1972. Quinacrine, a chemical stain specific for deoxyadenylate‐deoxythymilate rich regions in DNA. Proc. Natl. Acad. Sci. U.S.A. 69:629‐632.
   Weisblum, B. 1973. Fluorescent probes of chromosomal DNA structure: Three classes of acridines. Cold Spring Harbor Symp. Quant. Biol. 38:441‐449.
Key Reference for Q‐Banding
   Latt, S.A. 1976. Optical studies of metaphase chromosome organization. Annu. Rev. Biophys. Bioeng. 5:1‐37.
  During the 1970s, a number of review articles appeared that attempted to summarize all of the new developments in the field of cytogenetics. This article, with a 398‐item reference list, covers a variety of banding methods with detailed explanations of their underlying biochemistry. This review discusses Q‐banding as well as C‐banding, G‐banding, R‐banding, and the use of BrdU incorporation for chromosome labeling.
Literature Cited for G‐Banding
   Comings, D.E. 1978. Mechanisms of chromosome banding and implications for chromosome structure. Annu. Rev. Genet. 12:25‐46.
   Drets, M.E. and Shaw, M.W. 1971. Specific banding patterns of human chromosomes. Proc. Natl. Acad. Sci. U.S.A. 68:2073‐2077.
   Seabright, M. 1971. A rapid banding technique for human chromosomes. Lancet II:971‐972.
   Sumner, A.T. 1990. Chromosome Banding. Unwin Hyman, London.
   Sumner, A.T., Evans, H.J., and Buckland, R.A. 1971. New techniques for distinguishing between human chromosomes. Nature New Biol. 232:31‐32.
   van Duijn, P., van Prooijen‐Knegt, A.C., and van der Ploeg, M. 1985. Involvement of nucleosomes in Giemsa staining of chromosomes. A new hypothesis on the banding mechanism. Histochemistry 82:363‐376.
   Wang, H.C. and Federoff, S. 1972. Banding human chromosomes treated with trypsin. Nature New Biol. 235:52‐53.
Key References for G‐Banding
   Comings, 1978. See above.
  Review article with good discussions of possible underlying biochemical mechanisms of G‐banding in relationship to chromosome structure.
   Comings, D.E., Avelino, E., Harris, D.C., and Wyandt, H.E. 1973. The mechanism of C‐ and G‐banding of chromosomes. Exp. Cell Res. 77:469‐493.
  Good discussions of possible mechanisms of G‐banding.
   Sumner, 1990. See above.
  Effects of the pretreatments and function of the Giemsa stain.
   Sumner, A.T. and Evans, H.J. 1973. Mechanism involved in the banding of chromosomes with quinicrine and Giemsa. II. The interaction of the dyes with the chromosomal components. Exp. Cell Res. 81:223‐236.
  Presents a complex analysis of various steps in the G‐banding process and how these steps may be influenced by nucleosome structure of chromosomes.
   van Duijn, et al. 1985. See above.
Literature Cited for R‐Banding
   Latt, S.A., Jurgens, L.A., Matthews, D.J., Gustashaw, K.M., and Sahar, E. 1980. Energy transfer‐enhanced chromosome banding. Cancer Genet. Cytogenet. 1:187‐196.
   Sahar, E. and Latt, S.A. 1978. Enhancement of banding patterns in human metaphase chromosomes by energy transfer. Proc. Natl. Acad. Sci. U.S.A. 75:5650‐5654.
   Schweitzer, D. 1981. Counterstain enhanced chromosome banding. Hum. Genet. 57:1‐14.
   Sehested, J. 1974. A simple method of R‐banding of human chromosomes, showing a pH dependent connection between R‐ and G‐bands. Humangenetik 21:55‐58.
   Sumner, A.T. 1990. Chromosome Banding. Unwin Hyman, London.
Key References for R‐Banding
   Latt et al., 1980. See above.
  These two articles review the mechanisms of double‐dye staining.
   Schweitzer et al. See above.
Literature Cited for Replication Banding
   Gratzner, H., Ingram, D., Lief, R.C., and Castro, A. 1975. The use of antibody specific for bromodeoxyuridine for immunofluorescent determination of DNA replication in single cells and chromosomes. Exp. Cell Res. 95:88‐94.
   Korenberg, J. and Freedlander, E. 1974. Giemsa technique for detection of sister chromatid exchanges. Chromosoma 48:355‐360.
   Latt, S.A. 1973. Microfluorometric detection of deoxyribonucleic acid replication in human metaphase chromosomes. Proc. Natl. Acad. Sci. U.S.A. 70:3395‐3399.
   Latt, S.A., Willard, H.F., and Gerald, P.S. 1976. BrdU‐33258 Hoechst analysis of DNA replication in human lymphocytes with supernumerary or structurally abnormal X chromosomes. Chromosoma 57:135‐153.
   Lemieux, N., Dutrillaux, B., and Viegas‐Pequignot, E. 1992. A simple method for simultaneous R‐ or G‐banding and fluorescence in situ hybridization of small single‐copy genes. Cytogenet. Cell Genet. 59:311‐312.
   Perry, P. and Wolff, S. 1974. New Giemsa method for the differential staining of sister chromatids. Nature 251:156‐158.
Key Reference for Replication Banding
   Latt, S.A. 1976. Optical studies of metaphase chromosome organization. Annu. Rev. Biophys. Bioeng. 5:1‐37.
  An excellent review that covers the principles of replication banding.
Literature Cited for C‐Banding
   Comings, D.E., Avelino, E., Okada, T.A., and Wyandt, H.E. 1973. The mechanism of C‐ and G‐banding of chromosomes. Exp. Cell Res. 77:469‐493.
   Holmquist, G. 1979. The mechanism of C‐banding: Depurination and β‐elimination. Chromosoma 72:203‐224.
   Pardue, M.L. and Gall, J.G. 1970. Chromosomal localization of mouse satellite DNA. Science 168:1356‐1358.
   Pathak, S. and Arrighi, F.E. 1973. Loss of DNA following C‐banding procedures. Cytogenet. Cell Genet. 12:414‐422.
Key References for C‐Banding
   Sumner, A.T. 1990. Chromosome Banding. Unwin Hyman, London.
  These references provide detailed discussion of the mechanism of C‐banding.
   Holmquist, G. 1979. See above.
Literature Cited for NOR Staining
   Goodpasture, C. and Bloom, S.E. 1975. Visualization of nucleolus organizer regions in mammalian chromosomes using silver staining. Chromosoma 53:37‐50.
   Goodpasture, C., Bloom, S.E., Hsu, T.C., and Arrighi, F.E. 1976. Human nucleolus organizers: The satellites or the stalks. Am. J. Hum. Genet. 28:559‐566.
   Henderson, A.S., Warburton, D., and Atwood, K.C. 1972. Location of ribosomal DNA in the human chromosome complement. Proc. Natl. Acad. Sci. U.S.A. 69:3394‐3398.
   Howell, W.M. and Black, D.A. 1980. Controlled silver‐staining of nucleolus organizer regions with a protective colloidal developer: A 1‐step method. Experientia 36:1014‐1015.
   Howell, W.M., Denton, T.E., and Diamand, J.R. 1975. Differential staining of the satellite regions of human acrocentric chromosomes. Experientia 31:260‐262.
   Mikelsaar, A‐V., Schwarzacher, H.G., Schnedl, W., and Wagenbichler, P. 1977. Inheritance of Ag‐stainability of nucleolus organizer regions. Hum. Genet. 38:183‐188.
   Miller, D.A., Dev, V.G., Tantravahi, R., and Miller, O.J. 1976. Suppression of human nucleolus organizer activity in mouse‐human hybrid cells. Exp. Cell Res. 101:235‐243.
   Miller, O.J., Miller, D.A., Dev, V.G., Tantravahi, R., and Croce, C.M. 1976. Expression of human somatic cell hybrids. Proc. Natl. Acad. Sci. U.S.A. 73:4531‐4535.
Key Reference for NOR Staining
   Sumner, A.T. 1990. Chromosome Banding. Unwin Hyman, London.
  Contains a good discussion of the methodology and basis of silver staining.
Literature Cited for Distamycin/DAPI Staining
   Buckton, K.E., Spowart, G., Newton, M.S., and Evans, H.J. 1985. Forty‐four probands with an additional “marker” chromosome. Hum. Genet. 69:353‐370.
   Coll, M., Frederick, C.A., Wang, A.H‐J., and Rich, A. 1987. A bifurcated hydrogen bonded conformation in the d(A‐T) base pairs of the DNA dodecamer d(CGCAAATTTGCG) and its complex with distamycin. Proc. Natl. Acad. Sci. U.S.A. 84:8385‐8389.
   Miller, O.J., Schnedl, W., Allen, J., and Erlanger, B.F. 1974. 5‐Methylcytosine localized in mammalian constitutive heterochromatin. Nature 251:636‐637.
   Sahar, E. and Latt, S.A. 1980. Energy transfer and binding competition between dyes used to enhance staining differentiation in metaphase chromosomes. Chromosoma 79:1‐28.
  Schreck, R.R., Breg, W.R., Erlanger, B.F., and Miller, O.J. 1977. Preferential derivation of abnormal human G‐group‐like chromosomes from chromosome 15. Hum. Genet. 36:1‐12.
   Schweizer, D., Ambros, P., and Andrle, M. 1978. Modification of DAPI banding on human chromosomes by prestaining with a DNA‐binding oligopeptide antibiotic, distamycin A. Exp. Cell Res. 111:327‐332.
Key References for Distamycin/DAPI Staining
   Sahar and Latt, 1980. See above.
  Two excellent articles explaining dye interactions that lead to chromosome banding, including those between distamycin A and DAPI.
   Schweizer, D. 1981. Counterstain‐enhanced chromosome banding. Hum. Genet. 57:1‐14.
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