Table 1. DNA Content of Cell Nuclei DNA Content (Synonym) Cell Type or Source pg/nucleus Reference Mammalia

Man Brain cortex 7.1 31 Stomach 5.36 49 Liver 5.8–6.3 87 5.64–22.1 49 10.83 18 Pancreas 5.18 49 Cecum 5.16 49 Lung 6.04 49 WI-38 (embryonic lung fibroblast) 7.94 11 Leukocytes 6.85 52 Normal 6.92 56 11.06 30 Blood 7.63 18 Bone marrow 9.41 18 Leukemic 3.8 30 Bone marrow Lymphocytic leukemia 7.37 54 Granulocytic leukemia 8.88 54 Lymphocytes 5.06 49 5.61 82 6.50 55 Kidney 6.34 49 8.99 18 Sperm 1.15–3.50 57 2.43 45 2.44 49 2.7–3.4 72 3.40 18 Infertile 2.0 45 Prostate 9.43 18 Breast 6.50 49 Amnion, primary cell 18.8 29 HeLa (human cervix carcinoma) 14.4 13, 62 15.8 51 17 50 Suprarenal gland 9.1 43 Skin 5.60 49 Buffalo Sperm 4.90 22 Cat Brain cortex 7.1 31 Cattle Liver 5.9–6.2 47 5.9–10.5 48 6.1–6.4 86 6.22–8.4 58 6.4 87 Continued

1 Table 1. Continued Species (Synonym) Cell Type or Source DNA Content Reference pg/nucleus 6.5 70 6.8 46 Tetraploid 19.1 48 Pancreas 6.9 86, 87 Leukocytes 6.93–6.98 52 Lymphocytes 6.4 72 Spleen 7.0 46 Lymph node 7.03 58 Kidney 5.9 86 6.0 87 6.25–6.81 58 Sperm 2.8–3.3 47, 48 2.82 58 2.95 75 3.0 81 3.04 45 3.2 72, 89 3.24 71 3.3 86 3.9 91 4.44 21 Infertile 2.16 45 Calf Thymus 5.7–6.4 48 6.4 89 6.4–6.8 86 6.6 87 6.66 23 6.8 41 6.9 81 7.15 58 7.9 24 Dog Brain cortex 6.5 31 Liver 5.0–5.5 87 7.2 4 Leukocytes 6.73 52 Splenic lymphocytes 6.39 55 Kidney 5.3 87 Guinea pig Liver 5.9 86, 87 Horse Liver 5.8 87 Mouse Liver 6.0 87 6.6 10 12.3 43 Tetraploid 12.6 81 Octaploid 25.1 81 Hexadecaploid 48.8 81 Leukocytes 5.52 95 Transplant leukemia, AK4 7.4 61 9421 7.5 61 Lymphocytes 5.31 55 Continued

2 Table 1. Continued Species (Synonym) Cell Type or Source DNA Content Reference pg/nucleus 5.64 82 6.6 61 Granulocytic leukemia, C 1498 7.48 55 Spleen 6.5 65 8.3 37 Spontaneous leukemia 7.3 61 Thymus 7.9 94 8.78 73 Kidney 5.0 87 Fibroblasts, embryonic 5.1 7 LP-3 24.3 60 Carcinoma Krebs 7.0 38 Lettre-Ehrlich, hyperdiploid 11.1 37 Ehrlich, hypotetraploid 18.0 37 Ehrlich ascites cells 12.9–14.0 47 14.2 38 20.0 39 Ascites lymphoma, L1 32.2 73 L2 10.9 73 Cloudman S91 melanoma, tetraploid 16.6 37 6C3HED Lymphoma, diploid 9.8 37 6C3HED-DBA2 Lymphoma, hypotetraploid 15.3 37 DBA Ascites lymphoma 6.7 47 E9514A Lymphoma, diploid 7.9 37 Brush-tailed possum1 Unspecified 6.0 69 Rabbit Liver 5.3 87 7.23 16 Lymphocytes 5.75 72 Kidney, KD cell 8.7 51 Sperm 3.1 91 3.25 72 Rat Submaxillary gland 10.5 43 Salivary gland 6.85 78 Liver 5.9–9.4 15 6.0–8.6 48 6.9 93 7.8 20 7.98–9.31 78 8.2 46 10.8 2 11.5–13.0 43 JTC-25.P3 22.5 60 Pancreas 7.41 78 10.35 43 Small intestine 7.50 78 Lung 6.55 78 Leukocytes 6.1 81 6.75 78 Continued

3 Table 1. Continued Species (Synonym) Cell Type or Source DNA Content Reference pg/nucleus Lymphocytes 5.58 82 6.05 72 Bone marrow 6.90 78 Spleen 6.0–6.5 15 6.92 78 Thymus 6.1–6.3 15 6.72 23 7.2 78 Kidney 5.5 46 5.7 81 6.82 78 9.25 43 9.75 42 13.4 63 Spermatocytes, primary 15.2 24 Sperm 3.11 72 3.9 24 Sheep Liver 5.4–6.1 87 5.7 84 Leukocytes 6.83 52 Sperm 2.57 22 2.9 1, 91 Swine Liver 4.7–5.0 86 5.0 87 Leukocytes 6.85 52 Kidney 5.2 86, 87 Sperm 3.44 35

Aves

Agelaius phoeniceus Liver 3.7 5 Ardea herodias Liver 4.1 5 Cyanocitta cristata Liver 3.6 5 Dendroica coronata Liver 3.8 5 D. dominica Liver 3.9 5 D. palmarum Liver 4.4 5 Gallus gallus Erythrocytes 1.43 21 Sperm 0.73 21 (G. domesticus)2 Liver 2.39 58 2.45 70 2.52–2.77 17 3.5 5 Pancreas 2.51–2.83 16 Heart 2.67–2.75 16 Erythrocytes 1.70 41 2.2 89 2.34 58 2.4 83 Continued

4 Table 1. Continued Species (Synonym) Cell Type or Source DNA Content Reference pg/nucleus 2.5 81 2.54 66 2.62 16 2.63–2.70 17 3.1 91 3.30 72 Spleen 2.60–2.72 16 Kidney 2.38–2.57 16 Sperm 0.95 74 1.26 58 1.50 72 Fibroblasts 2.3 81 Embryo 1.4 91 Larus atricilla Liver 4.0 5 Melopsittacus undulatus Liver 3.3 5 Numida meleagris3 Diploid 2.27 85 Parus carolimemsis Liver 4.1 5 Pelecanus occidentalis Liver 3.9 5 Phalacrocorax auritus Liver 4.0 5 Pipilo erythrophthalmus Liver 3.6 5 Podiceps auritus Liver 4.2 5 Polioptila caerulea Liver 3.8 5 Regulus calendula Liver 3.7 5 Sayornis phoebe Liver 4.0 5 Serinus canarius Liver 4.1 5 Sialia sialis Liver 4.0 5 Sturnella magna Liver 4.1 5 Sturnus vulgaris Liver 3.6 5 Thalasseus sandvicensis Liver 3.9 5 Toxostoma rufum Liver 3.5 5 Duck Liver 2.1 87 Erythrocytes 2.3 85 2.65 85 Goose Diploid 1.9 88 2.92 85 Pheasant Diploid 1.7 88 Pigeon Diploid 2.0 88 Sparrow Diploid 1.9 88 Turkey Diploid 1.9 88

Reptilia

Chelonia mydas Liver 4.6–5.1 70 5.12 58 Erythrocytes 5.27 58 Alligator Erythrocytes 4.98 85 Black racer snake Erythrocytes 2.85 85 Pilot snake Erythrocytes 4.28 85 Continued

5 Table 1. Continued Species (Synonym) Cell Type or Source DNA Content Reference pg/nucleus Water snake Erythrocytes 5.02 85 Snapping turtle Erythrocytes 4.97 85 Wood turtle Erythrocytes 4.92 85

Amphibia

Alytes obstetricans Liver 21.0 14 Ambystoma mexicanum Liver 77.0 14 (Siredon mexicanum) Liver 10.7 24 Erythrocytes 6.88 23 11.2 24 Spermatocytes, primary 18.0 24 Sperm 3.36; 4.8 23, 24 Epithelium 10.9 24 A. tigrinum Liver 85.3 14 Amphiuma Erythrocytes 168 85 A. means Liver 192.8 14 Bombina bombina Liver 27.4 14 Bufo alvarius Erythrocytes 11.3 3 B. americanus Liver 10.8 14 Erythrocytes 12.7 3 B. bufo Liver 14.2 14 Erythrocytes 14.6 3 B. calamita Liver 9.2 14 Erythrocytes 10.7 3 B. fowleri Liver 12.6 14 Erythrocytes 14.2 3 B. marinus Liver 9.7 14 Erythrocytes 11.3 3 B. quercicus Erythrocytes 9.8 3 B. regularis Erythrocytes 8.9 3 B. retiformis Erythrocytes 13.4 3 B. terrestris Erythrocytes 11.1 3 B. valliceps Erythrocytes 9.2 3 B. viridis Erythrocytes 11.4 3 12.11 72 Sperm 5.4 72 Desmognathus fuscus Liver 36.0 14 Eurycea bislineata Liver 47.0 14 Hyla septentrionalis Liver 4.4 14 H. squirella Liver 10.4 14 Necturus Erythrocytes 48.4 85 N. lewisi Liver 270.0 14 N. maculosus Liver 196.8 14 N. punctatus Liver 264.0 14 Notophthalmus viridescens Liver 93.2 14 (Triturus viridescens) Diploid 98 76 Proteus anguinus Liver 105.1 14 Continued

6 Table 1. Continued Species (Synonym) Cell Type or Source DNA Content Reference pg/nucleus Rana catesbeiana Liver 15.2 14 R. esculenta Liver 14.1 14 Erythrocytes 14.6 53 Sperm 6.9 53 R. pipiens Liver 7.4–8.3 27 10.4 77 15.0 14 15.7 58 15.76 25 Erythrocytes 15.0 19, 58 Sperm 6.48 25 Ova 13.27 25 Embryo, late 13–16 77 14 9 R. temporatria Liver 8.8 14 Erythrocytes 8.2 91 8.9 72 (R. platyrrhina) Sperm 11–13 34 Scaphiopus bombifrons Liver 2.4 14 Triturus alpestris Liver 75.2 14 T. cristatus Liver 52.5 14 Erythrocytes 45.0 91 T. vulgaris Liver 71.0 14 Xenopus laevis Diploid 8.4 9 Liver 7.5 14 Erythrocytes 6.0 92 6.3 19 Toad Erythrocytes 7.33 58 Sperm 3.70 58

Dipnoi

Protopterus Erythrocytes 100 85

Crossopterygii

Latimeria chalumnae Erythrocytes 5.6 90 13.2 79

Osteichthyes

Acanthurus chirurgus Erythrocytes 1.38 59 (A. hepatus) Acipenser stellatus Sperm 2.12–2.5 36 A. sturio Erythrocytes 3.2 59 Aequidens portalegrensis Haploid 1.2 32 Alosa pseudoharengus Haploid 1.4 32 (Pomolobus pseudoharengus) Continued

7 Table 1. Continued Species (Synonym) Cell Type or Source DNA Content Reference pg/nucleus A. sapidissima Liver 1.9; 2.01 58, 70 Erythrocytes 1.97 58, 59 Haploid 1.3 32 Sperm 0.91 58 Amia Erythrocytes 2.3 59 Anguilla anguilla Erythrocytes 1.9 85, 88 A. rostrata Haploid 1.4 32 Anostomus anostomus Haploid 1.4 32 Antennarius ocellatus Haploid 0.78 32 Aphyocharax rubropinnis Haploid 1.7 32 Apteronotus albifrons Haploid 0.71 32 (Sternarchus albifrons) Arius felis (Galeichthys felis) Haploid 2.4 32 Aulostomus maculatus Erythrocytes 1.39 59 Bagre marinus Haploid 2.4 32 Balistes sp. Haploid 0.72 32 B. capriscus Erythrocytes 1.07 59 Barbus barbus Erythrocytes 3.4 85, 89 Betta splendens Haploid 0.64 32 Brachydanio rerio Haploid 1.8 32 Calamus calamus Erythrocytes 2.24 59 Callichthys callichthys Haploid 1.7 32 Caranx hippos Haploid 0.72 32 C. latus Erythrocytes 1.21 59 Carassius auratus Gill 3.43 8 Haploid 2.0 32 Centropristis striata Haploid 1.2 32 Cephalopholis fulva Erythrocytes 1.93 59 Haploid 1.2 32 Chela mouhoti Haploid 1.6 32 Chilodus punctatus Haploid 1.6 32 Cichlasoma biocellatum Haploid 1.3 32 C. meeki Haploid 1.4 32 Colisa lalia Haploid 0.62 32 lavaretus Diploid 5.8 6 widegrini (C. fera) C. muksun aspius Diploid 6.1 6 (C. macrophthalmus) aeneus Haploid 4.4 32 C. julii Haploid 4.2 32 C. melanistius Haploid 4.1 32 C. punctatus Haploid 2.9 32 C. undulatus Haploid 3.0 32 Culaea inconstans Haploid 0.67 32 (Eucalia inconstans) Cyprinus carpio Liver 2.8–3.2 87 3.33 58, 70 Erythrocytes 3.2 85, 88, 89 3.49 58, 59 Continued

8 Table 1. Continued Species (Synonym) Cell Type or Source DNA Content Reference pg/nucleus Haploid 1.7 32 Sperm 1.60 89 1.64 58 Dallia pectoralis Gill 2.53 8 Danio malabaricus Haploid 2.2 32 Decapterus punctatus Erythrocytes 1.32 59 Dermatolepis inermis Haploid 1.2 32 Diplodus argenteus Erythrocytes 1.61 59 Echeneis naucrates Haploid 0.72 32 Engraulis mordax Haploid 1.9 32 Eopsetta jordani Haploid 0.75 32 Epinephelus guttatus Liver 1.9–2.1 70 Erythrocytes 2.09 59 Haploid 1.2 32 E. morio Haploid 1.3 32 E. striatus Erythrocytes 2.05 59 Haploid 1.3 32 Esox americanus americanus Gill 2.37 8 E. americanus vermiculatus Gill 2.25 8 E. lucius Gill 2.72 8 Erythrocytes 1.7 85, 88, 89 Sperm 0.85 89 E. masquinongy Gill 2.56 8 E. niger Gill 2.37 8 E. reicherti Gill 2.56 8 Eucinostomus gula Erythrocytes 0.94 59 Exodon paradoxus Haploid 1.7 32 Fundulus heteroclitus Haploid 1.5 32 Gambusia affinis holbrooki Haploid 0.86 33 Gasterosteus aculeatus Haploid 0.70 32 Geophagus jurupari Haploid 1.2 32 Gymnocorymbus ternetzi Haploid 2.1 32 Gyrinocheilus aymonieri Haploid 0.65 32 Haemulon flavolineatum Liver 1.2–1.3 70 Erythrocytes 1.33 59 H. sciurus Erythrocytes 1.20 59 Harengula Sperm 6.85 26 H. humeralis (H. sardinia) Erythrocytes 2.04 59 Hemigrammus caudovittatus Haploid 1.7 32 H. ocellifer Haploid 1.7 32 Hippocampus erectus Haploid 0.66 32 (H. hudsonius) Holocentrus ascensionis Erythrocytes 1.31 59 Hoplias malabaricus Haploid 1.4 32 Huso huso Sperm 2.12–2.5 36 Hypostomus plecostomus Haploid 2.1 32 Ictalurus nebulosus Erythrocytes 1.80 85 (Ameiurus nebulosus) Continued

9 Table 1. Continued Species (Synonym) Cell Type or Source DNA Content Reference pg/nucleus Lactophrys quadricornis Erythrocytes 1.91 59 Loricaria parva Haploid 1.6 32 Lutjanus campechanus Haploid 1.4 32 L. griseus Erythrocytes 2.1 59 Haploid 1.3 32 L. vivanus (L. hastingsi) Erythrocytes 2.1 59 Lyopsetta exilis Haploid 0.73 32 Macropodus cupanus dayi Haploid 0.61 32 M. opercularis Haploid 0.59 32 Mastacembelus maculatus Haploid 0.74 32 Melanotaenia fluviatilis Haploid 1.3 32 Metynnis roosevelti Haploid 1.7 32 Misgurnus fossilis Sperm 2.6 80 Moenkhausia’oligolepis Haploid 1.6 32 Monacanthus hispidus Haploid 0.68 32 (Stephanolepis hispidus) Mugil curema Erythrocytes 1.38 59 Mycteroperca tigris Erythrocytes 2.2 59 Haploid 1.3 32 M. venenosa (Trisotropis Erythrocytes 1.83 59 venenosus) Novumbra hubbsi Gill 2.08 8 Ocyurus chrysurus Liver 2.1–2.4 70 Erythrocytes 2.1 59 Haploid 1.3 32 Ogcocephalus nasutus Haploid 0.74 32 Opsanus beta Haploid 3.0 32 O. tau Haploid 2.8 32 Otocinclus affinis Haploid 2.1 32 Otophidium scrippsae Haploid 0.68 32 Pantodon buchholzi Haploid 0.77 32 Paralabrax nebulifer Haploid 1.3 32 Parophrys vetulus Haploid 0.65 32 Peprilus triacanthus Haploid 0.80 32 (Poronotus triacanthus) Perca fluviatilis Erythrocytes 2.0 85, 89 Petrometopon cruentatum Haploid 1.2 32 Poecilia formosa Diploid 1.3–1.8 68 1.6 67 P. latipinna Diploid 1.6–1.9 68 Haploid 0.96 33 P. latipunctata Diploid 1.7–1.8 68 P. mexicana Diploid 1.6–1.9 68 P. reticulata Diploid 1.53 12 P. sphenops Diploid 1.7 67 1.8–1.9 68 P. vittata Diploid 1.9 67 Poeciliopsis latidens Diploid 1.30 12 Continued

10 Table 1. Continued Species (Synonym) Cell Type or Source DNA Content Reference pg/nucleus P. lucida Diploid 1.37 12 P. monacha Diploid 1.34 12 P. occidentalis Diploid 1.35 12 Porichthys notatus Haploid 2.2 32 Pterophyllum eimekei Haploid 1.2 32 Rachycentron canadum Haploid 0.77 32 Rutilus rutilus (Gardonus Erythrocytes 1.9 85, 88, 89 rutilus) Salmo gairdneri (S. irideus) Erythrocytes 4.9 85, 89 5.0 6, 40 5.61 72 Sperm 2.45 40, 89 2.52 72 S. gairdneri4 Sperm 6.5 26 S. trutta5 Erythrocytes 5.79 58 Salvelinus fontinalis Haploid 3.5 32 S. fontinalis6 Sperm 6.7 26 Scarus croicensis Erythrocytes 2.58 59 S. guacamaia (Pseudoscarus Erythrocytes 2.5 59 guacamaia) S. vetula Haploid 1.9 32 Scatophagus argus Haploid 0.77 32 Seriola rivoliana (Zonichthys Erythrocytes 1.35 59 falcatus) Sparisoma chrysopterum Erythrocytes 2.45 59 (S. brachiale) S. viride Haploid 2.3 32 (S. abildgaardi) Haploid 1.9 32 Sphoeroides maculatus Haploid 0.50 32 Sphyraena barracuda Erythrocytes 1.37 59 Stigmatogobius sadanundio Haploid 1.4 32 (Gobius sadanundio) Syngnathus fuscus Haploid 0.66 32 Synodus foetens Haploid 1.2 32 S. lucioceps Haploid 1.2 32 Tetraodon fluviatilis Haploid 0.40 32 Tinca tinca Erythrocytes 1.7 85, 89 Sperm 0.85 89 Trachinotus falcatus Haploid 0.85 32 Trichogaster leeri Haploid 0.82 32 T. trichopterus Haploid 0.73 32 Trinectes maculatus Haploid 0.65 32 Tylosurus acus Liver 2.2–2.5 70 Erythrocytes 2.2 59 Umbra limi Gill 5.03 8 Haploid 2.7 32 U. pygmaea Gill 4.82 8 Xenocara dolichoptera Haploid 1.8 32 Continued

11 Table 1. Continued Species (Synonym) Cell Type or Source DNA Content Reference pg/nucleus Xiphophorus helleri Haploid 0.95 33 1.42 12 1.6 64 X. maculatus Haploid 0.95 33 Xystreurys liolepis Haploid 0.74 32 Albacore Erythrocytes 1.79–2.23 28 Bigeye tuna Erythrocytes 1.85–2.09 28 Southern bluefin tuna Erythrocytes 1.91–2.17 28 Yellowfin tuna Erythrocytes 2.08–2.39 28 Moray eel Haploid 2.5 32

Chondrichthyes

Carcharhinus longimanus Erythrocytes 6.67 59 (Carcharias longimanus) Carcharhinus obscurus Liver 5.5 70 (Carcharias obscurus) Erythrocytes 5.46 59

Agnatha

Petromyzon Erythrocytes 5 59

Ascidiacea

Ascidia atra Sperm 0.16 59 Ciona intestinalis Sperm 0.14 44 1Trichosurus vulpecula. 2Chicken in reference. 3Guinea hen in reference. 4Rainbow trout in reference. 5Brown trout in reference. 6Brook trout in reference. Contributor: Shapiro, Herman S.

References 11. Christofalo, V. J., and D. Kritchevsky. 1969. Med. Exp. 19:313–320. 1. A˚ berg, B., and M. Gillner. 1966. Acta Physiol. Scand. 12. Cimino, M. C. 1974. Chromosoma 47:297–307. 66:106–114. 13. Cocito, C., and I. Meulemans. 1962. Acta Clin. Belg. 2. Albert, S., et al. 1953. Science 117:551–553. 17:74–75. 3. Bachmann, K. 1970. Chromosoma 29:365–374. 14. Conger, A. D., and J. H. Clinton. 1973. Radiat. Res. 4. Bachmann, K. 1972. Ibid. 37:85–93. 54:69–101. 5. Bachmann, K., et al. 1972. Ibid. 37:405–416. 15. Cunningham, L., et al. 1950. J. Gen. Physiol. 6. Bargetzi, J.-P. 1958. Experientia 14:445–447. 34:59–63. 7. Bassleer, R. 1964. C. R. Soc. Biol. 158:384–386. 16. Davidson, J. N., and W.M. Mclndoe. 1949. Biochem. J. 45:xvi. 8. Beamish, R. J., et al. 1971. Chromosoma 34:436–447. 17. Davidson, J. N., et al. 1950. Ibid. 46:xl. 9. Bristow, D. A., and E. M. Deuchar. 1964. Exp. Cell Res. 35:580–589. 18. Davidson, J. N., et al. 1951. Lancet 260:1287–1290. 10. Brodskii, V. Ya. 1955. Dokl. Akad. Nauk SSSR 19. Dawid, I. B. 1965. J. Mol. Biol. 12:581–599. 102:357–360. 20. Dounce, A. L., et al. 1950. J. Gen. Physiol. 33:629–642.

12 21. Eapen, K. J., and M. M. R. Nasir. 1963. Indian Vet. J. 52. Mandel, P., et al. 1950. C. R. Acad. Sci. 231:1172–1174. 40:803. 53. Manunta, G., and A. Marongiu. 1958. Arch. Sci. Biol. 22. Eapen, K. J., and M. M. R. Nasir. 1964. Indian J. Dairy (Bologna) 42:407–414. Sci. 17:107–108. 54. Menton, M. L., and M. WilIms. 1953. Cancer Res. 23. Edstro¨m, J-E. 1964. Biochim. Biophys. Acta 13:733–736. 80:399–410. 55. Menton, M. L., et al. l953. Ibid. 13:729–732. 24. Edstro¨m, J-E., and J. Kawiak. 1961. J. Biophys. Bio- 56. Metais, P., and P. Mandel. 1950. C. R. Soc. Biol. chem. Cytol. 9:619–626. 144:277–279. 25. England, M. C., and D. T. Mayer. 1957. Exp. Cell Res. 57. Meyho¨fer, W., et al. 1960. Arch. Klin. Exp. Dermatol. 12:249–253. 209:637–642. 26. Felix, K., et al. 1956. Hoppe Seylers Z. Physiol. Chem. 58. Mirsky, A. E., and H. Ris. 1949. Nature (London) 303:140–152. 163:666–667. 27. Finamore, F. J., et al. 1960. Arch. Biochem. Biophys. 59. Mirksy, A. E., and H. Ris. 1951. J. Gen. Physiol. 88:10–16. 34:451–462. 28. Fujii, Y., and S. Higasa. 1963. Nankai-Ku Suisan Ken- 60. Miyagishi, A., and T. Andoh. 1973. Biochim. Biophys. kyusho Hokohu 17:9–15 (Chem. Abstr. 59:l5641c, Acta 299:507–515. 1963). 61. Mizen, N. A., and M. L. Petermann. 1952. Cancer Res. 29. Gaffney, E. V., and J. Fogh. 1970. Proc. Soc. Exp. Biol. 12:727–730. Med. 133:607–608. 62. Mu¨ller, D. 1969. Virchows Arch. B2:6–17. 30. Gradova, M. G., et al. 1964. Vopr. Gematol. Pediatr. 63. Ogawa, K. 1961. Tex. Rep. Biol. Med. 19:825–832. Leningrad Sb. (3):300–309. 64. Ohno, S. 1970. Evolution by Gene Duplication. 31. Heller, I. H., and K. A. C. Elliot. 1954. Can. J. Biochem. Springer-Verlag, New York. Physiol. 32:584–592. 65. Petermann, M. L., and R. M. Schneider. 1951. Cancer 32. Hinegardner, R. 1968. Am. Nat. 102:517–523. Res. 11:485–489. 33. Hinegardner, R., and D. E. Rosen. 1972. Ibid. 66. Rasch, E., et al. 1971. Chromosoma 33:1–18. 106:621–644. 67. Rasch, E. M., et al. 1965. J. Exp. Zool. 160:155–170. 34. Hoff-Jørgensen, E., and E. Zeuthen. 1952. Nature 68. Rasch, E. M., et al. 1970. Chromosoma 31:18–40. (London) 169-245–246. 69. Rendel, J. M. 1955. Nature (London) 176:829. 35. Ivanov, I. I., et al. 1969. Byull, Eksp. Biol. Med. 70. Ris, H., and A. E. Mirsky. 1949. J. Gen. Physiol. 67(4):46–49 (Chem. Abstr. 71 :28329d, 1969). 33:125–146. 36. Kafiani, K. A., et al. 1958. Biochemistry (USSR) 71. Roberts, B., and S. Roberts. 1972. Chromosoma 23:389–399. 39:83–91. 37. Kit, S. 1960. Arch. Biochem. Biophys. 87:330–336. 72. Sandritter, W., et al. 1960. Acta Histochem. 38. Klein, E., and G. Klein. 1950. Nature (London) 10:139–154. 166:832–833. 73. Shelton, E. 1954. J. Natl Cancer Inst. 15:49–61. 39. Klein, E., et al. 1950. Exp. Cell Res. 1:127–134. 74. Solomon, J. B. 1957. Biochim. Biophys. Acta 40. Knobloch, A., et al. 1957: Biochim. Biophys. Acta 24:584–591. 24:201–202. 75. Summerhil, W. R., Jr., and D. Olds. 1961. J. Dairy Sci. 41. Kurnick, N. B. 1950. Exp. Cell Res. 1:151–158. 44:548–551. 42. Kurnick, N. B. 1951. J. Exp. Med. 94:373–376. 76. Swift, H., 1958. In W. D. McElroy., and B. Glass. ed. 43. Laird, A. K. 1954. Exp. Cell Res. 6:30–44. The Chemical Basis of Development. Johns Hopkins 44. Laird, C. D. 1971. Chromosoma 32:378–406. Univ. Press, Baltimore. pp. 174–213. 77. Sze, L. C. 1953. J. Exp. Zool. 122:577–601. 45. Leuchtenberger, C., and R. Leuehtenberger. 1958. Hoppe Seylers Z. Physiol. Chem. 313:130–137. 78. Thomson, R. Y., et al. 1953. Biochem. J. 53:460–474. 46. Leuchtenberger, C., et al. 1951. Proc. Natl Acad. Sci. 79. Thomson, K. S., et al. 1973. Nature (London) 241:126. USA 37:33–38. 80. Timofeeva, M. Ya., and K. A. Kafiani. 1964. Biochem- 47. Leuchtenberger, C., et al. 1952. Cancer Res. istry (USSR) 29:96–100. 12:480–483. 81. Tonkelaar, E. M. den, and P. van Duijn. 1964. Histo- 48. Leuchtenberger, C., et al. 1952. Exp. Cell Res. chemie 4:1–9. 3:240–244. 82. Utsumi, K. 1960. Acta Med. Okayama 14:1–13. 49. Leuchtenberger, C., et al. 1954. Am. J. Pathol. 83. Vaughn, J. C., and R. D. Locy. 1969. J. Histochem Cyto- 30:65–85. chem. 17:591–600. 50. Lin, H. J., and E. Chargaff. 1964. Biochim. Biophys. 84. Vendrely, R. 1950. Bull. Soc. Calm. Biol. 32:427–442. Acta 91:691–694. 85. Vendrely, R. 1958. Ann. Inst. Pasteur 94:143–166. 51. Littlefield, J. W., and E. A. Gould. 1960. J. Biol. Chem. 86. Vendrely, R., and C. Vendrely. 1948. Experientia 235:1129–1133. 4:434–436.

13 87. Vendrely, R., and C. Vendrely. 1949. Ibid. 5:327– 91. Walker, P. M. B., and H. B. Yates. 1952. Proc. R. Soc. 329. London B140:274–299. 88. Vendrely, R., and C. Vendrely. 1950. C. R. Acad. Sci. 92. Wallace, H., and M. L. Birnstiel. 1966. Biochim. 230:788–790. Biophys. Acta 114:296–310. 89. Vendrely, R., and C. Vendrely. 1953. Nature (London) 93. Weber, M. 1960. Acta Med. Pol. 1:27–45. 172:30–31. 94. Weymouth, P. P., et al. 1955. J. Natl Cancer Institute 90. Vialli, M. 1957. Rend. Ist. Lomb. Sci. Lett. I, 981–990. 91:680–686 (Chem. Abstr. 53:4588g, 1959). 95. Whitfeld, P. R. 1953. Aust. J. Biol. Sci. 6:234–243.

14 Table 2. RNA, DNA, and Protein Content of Vertebrate Cell Nuclei Tissue Constituent Value Medium or Technique Reference Nervous System

Brain Rat RNA/DNA 1.21 7

Digestive System

Salivary gland Mouse RNA/DNA 0.51 Detergent 33 Protein/DNA 3.2 Detergent 33 Liver Man Protein 42–59% 10 Acidic 35–51% 10 Other 4.7–7.5% 10 Cattle Nucleic acid 27.5–30.7% dry wt 17 RNA 0.9–1.9% dry wt 17 DNA 6.4 pg 6, 37 Calf RNA 1.5% dry wt Nonaqueous 1 DNA 15% dry wt Nonaqueous 1 22% dry wt Nonaqueous followed by citrate 1 Protein 85% dry wt Nonaqueous 1 74% dry wt Nonaqueous followed by citrate 1 Protein/DNA 5.7 Nonaqueous 1 3.35 Nonaqueous followed by citrate 1 Horse RNA 1.5% dry wt Nonaqueous 1 DNA 12% dry wt Nonaqueous 1 22% dry wt Nonaqueous followed by citrate 1 Protein 88% dry wt Nonaqueous 1 74% dry wt Nonaqueous followed by citrate 1 Protein/DNA 7.3 Nonaqueous 1 3.3 Nonaqueous followed by citrate 1 Fasted DNA 18% dry wt Nonaqueous 1 27% dry wt Nonaqueous followed by citrate 1 Protein 82% dry wt Nonaqueous 1 72% dry wt Nonaqueous followed by citrate 1 15 Continued 16 Table 2. Continued Tissue Animal Constituent Value Medium or Technique Reference Protein/DNA 4.55 Nonaqueous 1 2.7 Nonaqueous followed by citrate 1 Mouse RNA 3.4% dry wt 2 DNA 27.0% dry wt 2 Protein 66.0% dry wt 2 Protein/DNA 2.07 Chaveau followed by 0.35 M NaCl 9 3.56 Chaveau followed by 0.265 M KC1 9 2.75 Chaveau followed by Mg, Ca 9 Rabbit Nucleic acid 26.2% dry wt 17 RNA 2.0% dry wt 17 RNA/DNA 0.17 21 Rat RNA 2.9–7.6% dry wt 12, 17, 27 3% dry wt Aqueous 35 DNA 15% dry wt Aqueous 35 20–22% dry wt Aqueous 31 RNA/DNA 0.16 8 0.18–0.21 31 0.19 35 0.2 26 0.24 32 0.25 15 0.27 28 0.29 21 0.32 30 0.44 25 0.112–0.116 Detergent 5 0.117 Aqueous 5 0.22 Aqueous 38 0.48 Aqueous 38 Protein 40% dry wt Aqueous 35 Protein/DNA 4.0 8 2.67 Aqueous 35 4.8 Aqueous 38 Continued Table 2. Continued Tissue Animal Constituent Value Medium or Technique Reference 9.7 Aqueous 38 2.81 0.25 M sucrose, Ca 11 3.22 0.44 M sucrose, Ca 11 4.90 0.44 M sucrose, pH 3.8 11 5.80 0.44 M sucrose, pH 5.8 11 2.62 2.2 M sucrose, Ca 11 4.92 2.2 M sucrose, pH 5.7 11 Histone/DNA 0.62 0.44 M sucrose, Ca 11 1.9 0.44 M sucrose, pH 3.8 11 1.85 0.44 M sucrose, pH 5.8 11 0.52 2.2 M sucrose, Ca 11 1.05 2.2 M sucrose, pH 5.7 11 Fowl Nucleic acid 30.3% dry wt 17 RNA 2.1% dry wt 17 Chicken RNA/DNA 0.08 Detergent 14 0.16 Aqueous 14 Intestine Chicken RNA/DNA 0.30 Detergent 14 0.46 Aqueous 14 Pancreas Cattle DNA 6.6 pg 6, 37 17% dry wt Nonaqueous 1 Protein 81.5% dry wt Nonaqueous 1 Protein/DNA 4.8 Nonaqueous 1 Horse DNA 22% dry wt Nonaqueous 1 Protein 76% dry wt Nonaqueous 1 Protein/DNA 3.45 Nonaqueous 1

Circulatory System

Heart Cattle DNA 30.0% dry wt 3 16.5% dry wt Nonaqueous 1 Protein 83.5% dry wt Nonaqueous 1 Protein/DNA 5.1 Nonaqueous 1 17 Continued 18 Table 2. Continued Tissue Animal Constituent Value Medium or Technique Reference Calf DNA 19% dry wt Nonaqueous 1 Protein 80% dry wt Nonaqueous 1 Protein/DNA 4.2 Nonaqueous 1 Erythrocytes Fowl Nucleic acid 33.9–38.1% dry wt 17 RNA 0.8% dry wt 17 RNA/DNA 0.10 21 Chicken DNA 25.5% dry wt Nonaqueous 1 Protein 74.5% dry wt Nonaqueous 1 Protein/DNA 2.9 Nonaqueous 1 Goose DNA phosphorus 2.34% dry wt Nonaqueous 1 Bone marrow Calf DNA 23.5% dry wt Nonaqueous 1 Protein 76.5% dry wt Nonaqueous 1 Protein/DNA 3.3 Nonaqueous 1 Spleen Cattle Nucleic acid 32.6–33.6% dry wt 17 RNA 0.7–1.1% dry wt 17 DNA 6.8 pg 6, 37 Thymus Calf Nucleic acid 32% total nitrogen 19 DNA 6.4 pg 6, 37 26% dry wt Nonaqueous 1 30% dry wt Nonaqueous followed by citrate 1 RNA/DNA 0.06 21 0.036–0.067 0.25 M sucrose, Ca 13 Protein 74% dry wt Nonaqueous 1 74% dry wt Nonaqueous followed by citrate 1 Acidic 14% total nitrogen 19 Basic 36% total nitrogen 19 Protein/DNA 1.0–1.2 0.25 M sucrose, Ca 22 3.0 0.25 M sucrose, Ca 13 2.85 Nonaqueous 1 2.33 Nonaqueous followed by citrate 1 Fowl Nucleic acid 34.2% dry wt 17 RNA 1.4% dry wt 17 Continued Table 2. Continued Tissue Animal Constituent Value Medium or Technique Reference Urogenital System

Kidney Calf DNA 17% dry wt Nonaqueous 1 25% dry wt Nonaqueous followed by citrate 1 Protein 79% dry wt Nonaqueous 1 73% dry wt Nonaqueous followed by citrate 1 Protein/DNA 4.6 Nonaqueous 1 2.9 Nonaqueous followed by citrate 1 Rabbit Nucleic acid 26.0% dry wt 17 RNA 1.2% dry wt 17 Rat DNA 6.72 pg 36 RNA/DNA 0.13 30 Chicken DNA phosphorus 1.45% dry wt Nonaqueous 1 Sperm Cod Nucleic acid 30% dry wt 17 RNA 0.3% dry wt 17 Herring Nucleic acid 38.8–59.0% dry wt 17 RNA 0–0.2% dry wt 17 Salmon Nucleic acid 60.2% dry wt 17 RNA 0.1% dry wt 17

Tissue Culture Cells

Fibroblasts Chinese hamster DNA 7 pg 20 RNA/DNA 0.47 20 Acid-soluble protein/DNA 2.06 20 Acid-insoluble protein/DNA 1.38 20 WI-38 diploid fibroblasts Man RNA/DNA 0.44 Detergent 29 Protein/DNA 3.53 Detergent 29 HeLa Man RNA/DNA 0.24 ...... 4 0.32 Detergent 14 0.35 Detergent 23 0.29 Citrate, pH 3.0 23

19 0.45 Citrate, pH 3.9 23 Continued 20 Table 2. Continued Tissue Animal Constituent Value Medium or Technique Reference Protein/DNA; 3.1 Detergent 34 9.2 Detergent 23 6.9 Citrate, pH 3.0 23 8.4 Citrate, pH 3.9 23 Histone/DNA 3.6 Citrate 24

Tumors

Ascites carcinoma (TA3) Mouse Protein/DNA 2.71 Detergent 16 4.28 Detergent, Pb 16 Histone/DNA 0.72 Detergent 16 1.65 Detergent, Pb 16 Residual protein1 /DNA 1.85 Detergent 16 2.18 Detergent, Pb 16 Ascites sarcoma (Ehrlich-Lettre´)2 Mouse Protein/DNA 4.42 Detergent, Pb 16 Histone/DNA 1.61 Detergent, Pb 16 Residual protein1/DNA 2.49 Detergent, Pb 16 Ascites sarcoma (Ehrlich-Lettre´)3 Mouse Protein/DNA 5.93 Detergent, Pb 16 Histone/DNA 2.67 Detergent, Pb 16 Residual protein1/DNA 3.05 Detergent, Pb 16 Morris hepatoma Rat Protein/DNA 3.85 Detergent, Pb 16 Histone/DNA 1.93 Detergent, Pb 16 Residual protein1/DNA 1.69 Detergent, Pb 16 Walker carcinoma Rat RNA/DNA 0.79 32 Protein/DNA 3.45 Detergent 16 4.88 Detergent, Pb 16 Histone/DNA 0.66 Detergent 16 1.55 Detergent, Pb 16 Residual protein1/DNA 2.58 Detergent 16 3.07 Detergent, Pb 16 1Protein remaining associated with DNA after extraction of histones. 2Less than 7 da incubation. 314-19 da incubation. Contributors: Stein, Gary S., and Stein, Janet L. References 20. McBride, O. W., and E. A. Peterson. 1970. J. Cell Biol. 47:132–139. 1. Allfrey, V., et al. 1952. J. Gen. Physiol. 35:529–558. 21. McIndoe, W. M., and J. N. Davidson. 1952. 2. Barnum, C. P., et al. 1950. Arch. Biochem. Biophys. Brit. J. Cancer 6:200–214. 25:376–383. 22. Mirsky, A. E. 1971. Proc. Natl Acad. Sci. USA, 3. Behrens, M. 1932. Hoppe Seylers Z. Physiol. Chem. 68:2945–2948. 209:59–74. 23. Munro, G. F., et al. 1969. Exp. Cell Res. 55:46–52. 4. Berkowitz, D. M., et al. 1969. J. Cell Biol. 42:851–855. 24. Munro, G. F., et al. 1970. Cancer Res. 30:379–386. 5. Blobel, G., and V. R. Potter. 1966. Science 25. Muntwyler, E., et al. 1950. J. Biol. Chem. 154:1662–1665. 184:181–190. 6. Boivin, A., et al. 1948. C. R. Acad. Sci. 226:1061–1063. 26. Pogo, A. O., et al. 1966. Proc. Natl Acad. Sci. USA 7. Brody, T. M., and J. A. Bain. 1952. J. Biol. Chem. 56:550–557. 195:685–696. 27. Price, J. M., et al. 1950. Cancer Res. 10:18–27. 8. Chauveau, J., et al. 1956. Exp. Cell Res. 11:317–321. 28. Rosenthal, O., et al. 1956. J. Biol. Chem. 223:469–478. 9. Comings, D. E., and L. O. Tack. 1973. Ibid. 82:175–191. 29. Rovera, G., and R. Baserga. 1971. J. Cell. Physiol. 77:201–212. 10. Debov, S. S. 1951. Chem. Abstr. 45:10374. 30. Schneider, W.C., and V.R. Potter. 1949. J. Biol. Chem. 11. Dounce, A. L., and R. Ickowicz. 1969. Arch. Biochem. 177:893–903. Biophys. 131:359–368. 12. Dounce, A. L., et al. 1950. J. Gen. Physiol. 33:629–642. 31. Siebert, G. 1967. Methods Cancer Res. 2:287–301. 13. Frenster, J. H., et al. 1960. Proc. Natl Acad. Sci. USA 32. Steele, W., and H. Busch. 1963. Cancer Res. 46:432–444. 23:1153–1163. 14. Haussler, M. R., et al. 1969. Exp. Cell Res. 58:234–242. 33. Stein, G. S., and R. Baserga. 1970. J. Biol. Chem. 245:6097–6105. 15. Hogeboom, G. H., et al. 1952. J. Biol. Chem. 196:111–120. 34. Stein, G. S., and T. W. Borun. 1972. J. Cell Biol. 16. Magliozzi, J., et al. 1971. Exp. Cell Res. 67:111–123. 52:292–307. 17. Mauritzen, C. M., et al. 1952. Proc. R. Soc. London 35. Taylor, C. W., et al. 1973. Exp. Cell Res. 82:215–226. B140:18–3l. 36. Thomson, R. Y., et al. 1953. Biochem. J. 18. Maxwell, E., and G. Ashwell. 1953. Arch. Biochem. 53:460–474. Biophys. 43:389–398. 37. Vendrely, C. 1952. Bull. Biol. Fr. Belg. 86:1–87. 19. Mayer, D. T., and A. Gulick. 1942. J. Biol. Chem. 38. Widnell, C. C., and J. R. Tata. 1964. Biochem. J. 146:433–440. 92:313–317.

21 22 Table 3. Volume and Dry Mass of Mammalian Cell Nuclei Part I. Nuclear Volume Animal Cell Site Cell Type Volume, m3 Remarks Reference Man Ganglion, sympathetic Type I 624(137–1663) 3rd & 4th lumbar segments 12 Type II 1150(310–3262) Lung Bronchial epithelium 235 21 Ciliated columnar 436 Alveolar epithelium 528 Testis Spermatogonia 197(34–688) Small, resting nucleus 20 Spermatocytes, early primary 103(62–152) From peripheral layer Ovary Primary follicle Granulosa 50 Small, resting follicle 9 Tertiary follicle Granulosa 100 Follicle diameter, 2 mm Theca 150 Ruptured follicle Theca (180–200) Nuclear volume similar to granulosa nuclei at this stage Gravid follicle Granulosa lutein 300 1st month of pregnancy Corpus luteum Theca lutein 135 1st month of pregnancy 90 5th month of pregnancy Uterus: endometrium Basalis stratum 96(49–244) Young endometrium; non-pregnant 11 Functionalis stratum 192(91–643) female Vaginal epithelium Squamous 528 Surface epithelium from smear 22 Tissue culture Amnion 2552 2–5 da culture 13 HeLa 1037 2–11 da culture Guinea pig Ovarian follicle Cumulus oophorus Normal nuclei 69 1 Antrum Pyknotic nuclei 20 From cell lining antrum of old follicle Mouse Lung: bronchial epithelium lining Columnar (16–32) 16 Kidney 103 Castrate; diploid, compared to 2n liver 4 Proximal tubule DNA Collecting duct 60 Diploid, compared to 2n liver DNA Ovarian follicle Oocytes, early primary 438 One layer flattened cells 2 Medium 1365 One layer cuboidal cells Large 6010 2–3 layers of cells Tumor Sarcoma 333 Translated tumor; pyknotic stages 14 Fresh arbitrarily selected Pyknosis I 125 Pyknosis II 48 Continued Table 3. Continued Animal Cell Site Cell Type Volume, m3 Remarks Reference Pyknosis III 22 Embryo 1432 2-cell stage 2 705 4-cell stage 589 4- to 7-cell stage 373 8- to 12-cell stage Tissue culture, skin Fibroblasts 148 17 Rat Cerebellar cortex Piriform neurocytes 377(258–458) Vermian area: anterior lobe 6 (Purkinje cells) Gastric mucosa Chief 44 15 Parietal 57 Small intestine: crypts Columnar cells, goblet cells, & 160 7 intestinal cells with acidiform granules (Paneth cells) Large intestine: crypts Columnar cells, goblet cells, & 80 7 intestinal cells with acidiform granules (Paneth cells) Pancreas: islets of Langerhans A cells 62(16–125) 5 B cells 88(13–167) Kidney 105 F proximal portion. C straight 8 Convoluted tubule tubule 14–23% larger than Straight tubule 102 convoluted tubule Uterine gland 77 Castrate. Basic volume, diploid, 3 compared to 4n liver DNA Suprarenal gland 18 Cortex Fasciculata 100 Medulla 140 Thyroid 68 Diploid, compared to 2n liver 4 DNA 1 42 months old Liver parenchyma 2n (diploid) 75(38–109) 10 4n 185(119–210) 8n 340(294–404) 14 months old Liver parenchyma 4n 268(195–320) 10 8n 545(380–761)

23 Values in parentheses are ranges. Contributor: Lung, Ben. References 11. Hintzsche, E. 1949. Gynaecologia 128:270–285. 12. Jonas, F.J. 1951. Z. Zellforsch. Mikrosk. Anat. 35:333–356. 1. Alfert, M. 1955. Biol. Bull. 109:1–12. 13. Lampert, F., et al. 1966. Histochemie 6:370–381. 2. Alfert, M. 1950. J. Cell Comp. Physiol. 36:381–409. 14. Leuchtenberger, C. 1950. Chromosoma 3:449–473. 3. Alfert, M., and H. A. Bern. 1951. Proc. Natl Acad. Sci. USA 37:202–205. 15. Leuchtenberger, C., and F. Schrader. 1951. Biol. Bull. 4. Alfert, M., and I. I. Geschwind. 1953. Ibid. 101:95–98. 39:991–999. 16. Leuchtenberger, C., et al. 1958. Cancer 11:490–506. 5. Arnold, A. 1951. Acta Anat. 12:396–428. 17. Mellors, R. C., et al. 1953. Ibid. 6:372–384. 6. Attardi, G. 1957. Exp. Cell Res., Suppl. 4:25–53. 18. Rather, L. J. 1951. J. Exp. Med. 93:573–586 7. Bergemann, T. 1952. Z. Mikrosk. Anat. Forsch. 19. Rather, L. J. 1958. Ergeb. Allg. Pathol. Pathol. Anal. 58:196–207. 38:127–199. 8. Deimling, O. von., et al. 1968. Histochemie 12:165– 20. Roosen-Runge, E. C., and F. D. Barlow. 1953. 174. Am. J. Anat. 93:143–166. 9. Dubner, R. 1952. Z. Mikrosk. Anat. Forsch. 58:147– 21. Sandritter, W., et al. 1958. Frankf. Z. Pathol. 195. 69:167–193. 10. Heizer, P. 1955. Chromosoma 7:281–327. 22. Sandritter, W., et al. 1963. Acta Cytol. 7:45–53.

24 Table 3. Part II. Nuclear Dry Mass Animal Cell Site Cell Type Dry Mass Pg Reference Man Buccal mucosa Parabasal 42.8 7 Intermediate 61.9 Rod-pyknotic 48.1 Pyknotic 34.5 Lung Basal 33.2 17 Bronchial epithelium Ciliated columnar 41.1 Oat cell carcinoma 34.5+19 19 Alveolar epithelium 47.5 17 Pleura Mesothelial 37.4 +16.2 19 Sperm 6.51 11 Vagina Normal superficial 43.0 16 Carcinoma 90.8 Tissue culture Amnion 166.3 6 HeLa 100.7 Child Blood Lymphocytes 48.2 5 Adult Blood Lymphocytes 50.31 3 Cattle (15.4–35.0)2 4 Calf Thymus Bull Sperm 7.1 8 Guinea pig Sperm 11.5(11–12) 13 Mouse Liver 2n (diploid) 45.6 2 4n 97.5 8n 190.5 Lung (16.5–19.0) 9 Testis Germinal cell; diploid; prophase 29(27–31) 15 Sperm 13.0(12–14) 13 Uterine cervix Parabasal 54.8 1 Intermediate; precornified 32.1 Superficial; cornified 16.8 Hyperplasia 74.8 Invasive cancer 108.5 Tissue culture Sarcoma 88 14 Continued 25 26

Table 3. Continued Animal Cell Site Cell Type Dry Mass Pg Reference Embryo Tissue culture Fibroblasts 74 14 Rabbit Cerebellar cortex Piriform neurocytes (Purkinje 364.0 10 cells) Liver Hepatocytes 44.4 12 Testis Spermatocytes 6.71 þ 0.90 18 Rat Liver Hepatocytes 79.4 12 Testis Spermatocytes 5.82 þ 0.51 18 1Determined by quantitative electron microscopy. 2Range of means. Values in parentheses are ranges. Contributor: Lung, Ben References 10. Lodin, Z., et al. 1967. Brain Res. 6:489–509. 11. Lung, B. 1974. Adv. Cell Mol. Biol. 3:73–132. 1. Broghamer, W. L., and W. M. Christopherson. 1961. 12. Mclndoe, W. M., and J. N. Davidson. 1952. Cancer 14:378–383. Br. J. Cancer 6:200–214. 2. Davies, H. G., et al. 1957. Exp. Cell Res., Suppl. 13. Mellors, R. C., and J. Hlinka. 1955. Exp. Cell Res. 4:136–149. 9:128–134. 3. Golomb, H. M., and G. F. Bahr. 1973. Chromosoma 14. Mellors, R. C., et al. 1953. Cancer 6:372–384. 46:233–245. 15. Mellors, R. C., et al. 1954. Ibid. 7:873–883. 4. Hale, A. J. 1957. Acta Histochem. 4:222–229. 16. Sandritter, W. 1958. Acta Cytol. 2:321–323. 5. Lampert, F. 1968. Virchows Arch. B 1:31–38. 17. Sandritter, W., et al. 1958. Frankf. Z. Pathol. 69:167– 6. Lampert, F., et al. 1966. Histochemie 6:370–381. 193. 7. Lee, L., et al. 1973. Acta Cytol. 17:214–219. 18. Sandritter, W., et al. 1958. Verh. Anat. Ges. 105:146– 8. Leuchtenberger, C., et al. 1956. Chromosoma 156. 8:73–86. 19. Sandritter, W., et al. 1960. Klin. Wochenschr. 38:590– 9. Leuchtenberger, C., et al. 1958. Cancer 11:490–506. 595.

27 28

Table 4. Macromolecules Characteristic of Cell Nuclei The examples given in this table are only the best defined and most insoluble of nuclear macromolecules. There are undoubtedly many other nuclear macromolecules that have eluded description because of technical difficulties of nuclear isolation. Many are rather weakly bound to nuclei and appear to be cytoplasmic because of elution from nuclei during isolation. Others may not have been stabilized sufficiently to assay reliably. Still others, such as “acidic proteins,” are well known and are poorly defined at present, and cannot be catalogued in a table. Enzyme Commission No. ¼ number assigned by the Commission on Biochemical Nomenclature [ref. 3]. Subspecies: Nomenclature used is that given in the literature cited; number followed by “S” ¼ sedimentation coefficient in Svedbergs. Data in brackets refer to the column heading in brackets.

Molecule (Synonym) Subspecies Molecular [Enzyme Commission No.] (Synonym) Location Weight daltons Method of Solubilization Reference Histones HI (Fl) Nucleus only 20,000 All subspecies: Dilute acid, e.g., 1 N HC1; or high 5 H2A (F2a2) Nucleus only 14,200 salt, e.g., 1 M NaCl H2B (F2b) Nucleus only 13,800 H3 (F3) Nucleus only 15,200 H4(F2al) Nucleus only 11,300 Ribosomal RNA’s: precursors & 45S Nucleolus only 4.1106 All subspecies extracted into aqueous phase of 12 products1 32S Nucleolus only 2.1106 emulsion of physiological saline and phenol 28S Nucleolus & cytoplasm 1.65106 and/or chloroform 20S Nucleolus only 0.95106 18S Predominantly cytoplasm 0.65106 5.5S Nucleolus & cytoplasm 0.04106 NMN adenylyltransferase [2.7.7.1] Nucleus, except in large Not easily solubilized 6, 10 neurons and in Acetabularia 6 RNA nucleotidyltransferase (RNA I (A) Nucleolus (4–5)10 All subspecies: 0.32 M (NH4)2SO4 plus 1, 8, 9 polymerase) [2.7.7.6] II (B) Nucleoplasm (4–5)106 sonication III (C) Nucleoplasm DNA nucleotidyltransferase (DNA 6.8S (a) Nucleus in non-aqueous 140,000; Many aqueous solutions 13 polymerase) [2.7.7.7] extraction; otherwise, 110,000 cytoplasmic 3.4S (b) Nucleus only 43,000 High salt, e.g., 1 M NaCl; pH, 8.0 2, 4, 7 Poly-(ADP-ribose) synthetase Nucleus only 78,000 Not easily solubilized 11 1 Mammalian cells. Contributor:Gu˙ rney, Theodore, Jr. References 7. Kier, H. M., et al. 1962. Nature (London) 196:752– 754. 1. Chambon, P.1975. Annu. Rev. Biochem. 44:613–638. 8. Schwartz, L. B., and R. G. Roeder. 1974. J. Biol. Chem. 2. Chang, L. M. S., et al. 1973. J. Mol. Biol. 74:1–8. 249:5898–5906 3. Commission on Biochemical Nomenclature. 1973. 9. Schwartz, L. B., et al. 1974. Ibid. 249:5889–5897. Enzyme Nomenclature (1972). American Elsevier, New York. 10. Siebert, G. 1972. Sub-Cell Biochem. 1:277–292. 4. Foster,D.N.,andT.Gurney,Jr.1974.J.CellBiol.63:103a. 11. Sugimura, T. 1973. Prog. Nucleic Acid Res. Mol. Biol. 5. Hnilica, L. S. 1972. The Structure and Biological Func- 13:127–151. tions of Histones. CRC Press, Cleveland. 12. Weinberg, R. A. 1973. Annu. Rev. Biochem. 6. Kato, T., and O. H. Lowry. 1973. J. Biol. Chem. 42:329–354. 248:2044–2048. 13. Weissbach, A. 1975. Cell 5:101–108.

29 30 Table 5. Isolation and Chemical Composition of Mammalian Cell Nuclei Methods for isolation of mammalian cell nucleoli [ref. 2 18] are dependent upon isolation of mammalian cell nuclei in aqueous media. The nuclei are usually prepared by homogenization in sucrose solutions [ref. 2] containing divalent metal ions (Ca2þ ,Mg2þ ,orMn2þ ), which are required to main- tain nucleolar integrity. (Detergent solutions have been used, but the nucleolar morphology is altered [ref. 17, 18]. A method employing nuclei homogenized in citric acid has been of some value for tissues from which satisfactory “sucrose-nuclei” have not been prepared.) Nucleoli and other nuclear structures are released by sonic oscillation [ref. 2], repeated compression-decompression [ref. 2, 7], or by enzymatic digestion with deoxy- ribonuclease I (DNase) [ref. 18]. The nucleoli are resistant to physical or enzymatic treatments because they are compact structures, have high mechanical strength, and are surrounded by a layer of perinucleolar chromatin. Differential centrifugation through sucrose gradients is used to purify the nucleoli. Several washings may be required to eliminate extranucleolar chromatin. Using these procedures, which can be adapted readily to particular mammalian cells, nucleoli have been prepared from liver [ref. 10], many cells in tissue culture [ref. 17, 18], various types of tumors [ref. 3, 4, 6], and brain [ref. l]. The criteria for purity include light and phase micro- scopic examination of the preparations for numbers and quality of stained (with Azure C) and unstained nucleoli, as well as a search for cytoplasmic and nucleoplasmic contaminants [ref. 16]. Electron microscopic analysis demonstrates the presence or absence of nuclear components, as well as microsomes, ribosomes, and membranes if present. The method of choice is the sucrose-Ca2þ isolation of nuclei, followed by sonication for nuclear disruption, because it is widely applicable and simple [ref. 2]. The DNase method [ref. 17] reduces DNA content, but is convenient for isolation of ribonucleoprotein particles. The combined citric acid-sonication method [ref. 15] results in loss of RNA nucleotidyltransferase(RNA polymerase) activity, but is very convenient when nuclear isolation is difficult. For detailed analyses of chemical constituents in terms of fractionation, consult references 2 and 14 for low molecular weight RNAs, references 2 and 5 for intermediate and high molecular weight RNAs, references 2 and 16 for enzymes, reference 12 for proteins, reference 11 for phosphoproteins, and reference 13 for preribosomal proteins [ref. 13]. Lipids and carbohy- drates were not included because of insufficient data. Chemical Constituent: W ¼ sum of weights of DNA, RNA, and protein. Data in brackets refer to the column heading in brackets.

Chemical Constituent % W [pg/nucleolus]

Cell Type or Tissue Animal Method of Isolation DNA RNA Protein Reference Liver Guinea pig Sonication 9.5 4.1 86.4 9 Rat Compression–decompression 8.7 [0.57] 5.9 [0.39] 85.4 [5.6] 7 3.0 [0.22] 7.5 [0.55] 89.5 [6.5] 4 Sonication 2.8 [0.38] 2.8 [0.38] 94.4 [12.8] 4 6-hours regeneration Rat Sonication 4.6 7.6 87.8 10 18-hours regeneration Rat Sonication 5.4 15.5 79.1 10 Tumors Man Sonication 6–331 67–94 3 Ascites Mouse Sonication 9.9 10.9 79.2 6 Continued Table 5. Continued Cell Type or Tissue Animal Method of Isolation Chemical Constituent % W [pg/nucleolus] Reference

DNA RNA Protein Hepatomas Novikoff Rat Compression–decompression 7.5 [2.15] 8.0 [2.30] 84.5 [24.3] 8 Sonication 13.5 13.5 73.0 15 Zajdela Rat Sonication 6 [1.6] 12 [3.0] 82 [23.0] 19 Walker Rat Compression–decompression 11.7 [1.6] 16.2 [2.2] 72.1 [9.8] 4 Sonication 6.7 [1.7] 7.5 [1.9] 85.8 [21.9] 4 1Value for sum of RNA þ DNA; RNA/DNA ¼ 0.27–0.56. Contributors: Choi, Yong C., and Busch, Harris 31 References 10. Muraxnatsu, M., and H. Busch. 1965. J. Biol. Chem. 240:3960–3966. 1. Banks, S. P., and T. C. Johnson. 1973. Biochhn. Bio- 11. Olson, M. O. J., et al. 1974. Ibid. 249:2823–2827. phys. Acta 294:450–460. 12. Orrick, L. R., et al. 1973. Proc. Natl Acad. Sci. USA 2. Busch, H., and K. Smetana. 1970. The Nucleolus. Aca- 70:1316–1320. demic Press, New York. 13. Prestayko, A. W., et al. 1974. FEBS Lett. 44(2):131–135. 3. Busch, H., et al. 1965. Cancer Res. 25:225–233. 14. Ro-Choi, T. S., and H. Busch. 1974. In H.. Busch. ed. 4. Busch, H., et al. 1966. Natl Cancer Inst. Monogr. The Cell Nucleus. Academic Press, New York. v. 3, 23:193–212. pp. 151–208. 5. Choi, Y. C. et al. 1974. In H. Busch. ed. The Cell 15. Ro-Choi, T. S., et al. 1973. Exp. Cell Res. 79:43–52. Nucleus. Academic Press, New York.v. 3, pp. 109–149. 16. Siebert, G., et al. 1966. J. Biol. Chem. 241:71–78. 6. Izawa, M., and K. Kawashima. 1967. Biochjm. Bio- phys. Acta 155:51–62. 17. Vesco, C., and S. Penman. 1968. Biochim. Biophys. 7. Liau, M. C. 1969. Adv. Enzyme Regul. 7:219–233. Acta 169:188–195. 8. Liau, M. C., et al. 1965. Proc. Natl Acad. Sci. USA 18. Zalta, J., and J-P. Zalta. 1973. Methods Cell Biol. 53:626–633. 6:317–324. 9. Maggio, R., et al. 1963. J. Cell Biol. 18:293–312. 19. Zalta, J., et al. 1971. J. Cell Biol. 51:563–568.

32 Table 6. Chromatin Components Relative to DNA Chromatin is defined as the diffuse interphase form of the chromosomes of eukaryote cells. This structure includes several macromolecular compo- nents in addition to DNA: (i) small amounts of RNA; (ii) histones, in mass about equal to the mass of DNA; and (iii) a variable amount of non-his-tone proteins. For a recent review of the subject, consult reference 28. Values for composition and optical proteins [ref. 15], as well as for template activity [ref. 7, 29], depend on the procedure employed for isolation of the chromatin. The first mandatory step for chromatin isolation is the preparation of highly purified nuclei to avoid cytoplasmic and membrane contamination of nucleoproteins [ref. 29]. Data are for different tissues of different species, and for tumors and isolated cell lines.

Source RNA Total Protein Histone Residual Protein

Cell Type or Tissue Organism & Specification DNA DNA DNA DNA Reference Vertebrates

Brain Cat 0.110 3.02 1.86 18 Cerebellum Swine 0.130 1.60 0.50 25 Liver Rat 0.043 1.67 1.00 0.67 19 Normal 0.10 2.2 32 Chick 0.025 2.61 24 Regenerating, 1 hour Rat 0.06 2.3 32 5 hours Rat 0.07 2.4 32 24 hours Rat 0.10 2.8 32 48 hours Rat 0.10 3.0 32 Intestinal mucosa Rat 0.120 3.70 11 Cardiac muscle Dog, 8–9 years old 0.036 1.84 27 Erythrocytes Chick 0.003 1.18 24 Thymus Calf 1.46 1.14 0.32 16 Kidney Chick 0.007 1.76 24 Prostate, ventral Rat 0.090 1.79 1.09 0.70 6 Mammary gland Rat 2 Not pregnant 0.110 1.31 0.74 Pregnant, 16–21 da 0.130 1.32 0.55 Lactating, 4 da 0.250 1.62 0.84 Hypophysis1 Swine 0.108 1.56 0.45 25

33 Continued 34

Table 6. Continued Source RNA Total Protein Histone Residual Protein Reference

Cell Type or Tissue Organism & Specification DNA DNA DNA DNA Embryo Chick, 11 da 0.151 1.17 0.30 14 SV40-transformed cells Mouse 0.050 2.49 1.01 1.48 26 Morris hepatoma Rat 0.120 2.00 1.11 0.90 1 5123 C 9618 A Rat 0.070 2.06 1.20 0.9 Ehrlich ascites cells Mouse 0.082 2.11 1.12 0.94 17 Ovary cells Chinese hamster 0.18 1.14 1.37 9 HeLa S-3 cells Man 0.050 1.80 0.70 1.08 3

Invertebrates

Salivary gland Drosophila 0.030 2.00 0.97 1.03 12 Midgut gland Drosophila 0.090 2.23 0.98 1.25 12 Sperm Spider crab 2.75 30 Sea urchin 0.040 1.02 0.13 21 Embryo, 0–2 hours Drosophila 0.060 1.88 0.60 1.28 8 6–18 hours Drosophila 0.060 1.98 0.79 1.19

Protozoa

Ciliate Tetrahymena pyriformis 0.071 1.41 0.93 0.48 10

Algae

Dinoflagellate 0.090 0.56 0.08 0.48 22 Continued Table 6. Continued Source RNA Total Protein Histone Residual Protein Reference

Cell Type or Tissue Organism & Specification DNA DNA DNA DNA Fungi

Spheroplasts Neurospora crassa 0.140 0.84 0.24 0.60 13 Yeast Saccharomyces cerevisiae 3.800 9.7 1.0 8.7 31 Plasmodium Physarum polycephalum2 0.3 10.2 1.10 9.1 20

Higher Plants

Root 23 Cortex Rhoeo discolor 2.30 13.3 Meristematic zone Rhoeo discolor 2.50 5.1 Central cylinder Rhoeo discolor 2.50 12.3 Seedling Pea 0.50 2.67 1.13 1.54 4 Embryo, 1 cm long Pea 0.013 1.03 5 1Synonym: Pituitary. 2Slime mold. Contributor: Vidali, Giorgio. 35 References 15. Johnson, R. S., et al. 1972. Biochemistry 11:4347–4358. 16. Kielman, L., and R. C. C. Huang. 1971. J. Mol. Biol. 1. Arnold, E. A., et al. 1973. Cancer Res. 33:1169–1176. 55:503–521. 2. Barker, K. L., and T.M. Ludwick. 1974. Proc. Soc. Exp. 17. Kostraba, N. C., et al. 1975. J. Biol. Chem. 250:1548– Biol. Med. 145:1325–1328. 1555. 3. Bhorjee, J. S., and T. Pederson. 1973. Biochemistry 18. Lu, C. Y., and H. Koenig. 1973. FEBS Lett. 34:48–54. 12:2766–2773. 19. Marushige, K., and J. Bonner. 1966. J. Mol. Biol. 4. Birnstiel, M. L., and B. B. Hyde. 1963. J. Cell Biol. 15:160–174. 18:41–50. 20. Mohberg, J., and H. P. Rusch. 1970. Arch. Biochem. 5. Bonner, J., and R-C. Huang. 1963. J. Mol. Biol. Biophys. 138:418–432. 6:169–174. 21. Ozaki, H. 1971. Dev. Biol. 26:209–219. 6. Couch, R. M., and K. M. Anderson. 1973. Biochemis- 22. Rizzo, P. J., and L. D. Nooden. 1972. Science try 12:3114–3121. 176:796–797. 7. de Pomerai, D. I., et al. 1974. Eur. J. Biochem. 23. Ruch, F., and A. Rosselet. 1970. Exp. Cell Res. 62:219– 46:461–471. 227. 8. Elgin, S. C. R., and L. F. Hood. 1973. Biochemistry 24. Seligy, V., and M. Miyagi. 1969. Ibid. 58:27–34. 12:4984–4991. 25. Shaw, L. M. J., and R. C. C. Huang. 1970. Biochemistry 9. Gerner, E. W., and R. M. Humphrey. 1973. Biochim. 9:4530–4542. Biophys. Acta 331:117–127 26. Shih, T. Y., et al. 1973. Proc. Natl. Acad Sci. USA 10. Hamana, K., and K. Iwai. 1971. J. Biochem.(Tokyo) 70:3506–3510. 69:1097–1111. 27. Shirey, T. L., and H. Sobel. 1972. Exp. Gerontol. 11. Haussler, M. R., et al. 1968. J. Biol. Chem. 243:4055– 7:15–29. 4064. 28. Simpson, R. T. 1973. Adv. Enzymol. 38:41–108. 12. Helmsing, P. J., and O. van Eupen. 1973. Biochim. 29. Tata, J. R., et al. 1972. J. Mol. Biol. 67:231–246. Biophys. Acta 308:154–160. 30. Vaughn, J. C., and G. W. Hinsch. 1972. J. Cell Sci. 13. Hsiang, M. W., and R. D. Cole. 1973. J. Biol. Chem. 11:131–152. 248:2007–2013. 31. Wintersberger, U., et al. 1973. Eur. J. Biochem. 14. Huang, R. C. C., and P. C. Huang. 1969. J. Mol. Biol. 33:123–130. 39:365–378. 32. Yeoman, L. C., et al. 1975. Cancer Res. 35:1249–1255.

36 Table 7. Chemical and Enzymic Composition of the Nuclear Envelope Techniqueshave been developed specifically for isolation of the nuclear envelope from rat liver [ref. 1, 5, 10, 12, 17, 19, 23], swine liver [ref. 10], bovine liver [ref. 3, 4], hen erythrocytes [ref. 24], rat and calf thymus [ref. 11, 18], and rat ventral prostate [ref. 20]. Additional methodological details have been published [ref. 2, 14, 22], and the structural and functional organization of the nuclear envelope has been the subject of several reviews [ref. 6–9,13,16,21]. Notwithstanding published procedures, it should be emphasized that careful microscopic examination of both the nuclear preparation and the nuclear envelope fractions is essential in establishing that the procedure is being successfully employed. In most cases, fractionation methods have been monitored by electron microscopy, and suitable analytical measurements (chemical and biochemical) have been applied to permit eval- uation of the degree of contamination byothercell membrane systems. The data below are for nuclear envelope preparations which meet these criteria. Part I. General Composition Cell Source [Buoyant Density, g/ml] Constituent Value Reference Liver Bovine [1.05-1.23] Phospholipid 22.6 4 Protein 70.4 Ribonucleic acid 5.8 Deoxyribonucleic acid 1.1 Rat [1.16] Phospholipid 27–50 23 Sialic acid(s) 0.09 Protein 25 Ribonucleic acid 2.1 Deoxyribonucleic acid 0.2 [1.16–1.18]1,2 Phospholipid 30.1 14 Cholesterol 2.3 Neutral sugars3 2.7 Hexosamines 0.24 Sialic acid(s) 0.06 Protein 61.3 Ribonucleic acid 3.3 Deoxyribonucleic acid 0 [1.17–1.18] Phospholipid 23 19 Protein 73 Ribonucleic acid 3 Deoxyribonucleic acid 0.6 [1.18–1.19] Phospholipid 38 5 Protein 52 Ribonucleic acid 10 Deoxyribonucleic acid Trace [1.18–1.20]1,5 Phospholipid 25.0 14 Cholesterol 1.0 Neutral sugars3 3.7 Hexosamines4 0.21 Sialic acid(s) 0.09 Protein 63.7 Ribonucleic acid 6.3 Deoxyribonucleic acid 0

Continued

37 Table 7. Continued Cell Source [Buoyant Density, g/ml] Constituent Value Reference [1.19] Phospholipid 27–50 23 Sialic acid(s) 0.09 Protein 47 Ribonucleic acid 3.3 Deoxyribonucleic acid 1.3 [1.215] Phospholipid 16.1 10 Other lipids6 2.8 Protein 75.5 Ribonucleic acid 3.6 Deoxyribonucleic acid 2.0 [1.27] Phospholipid 23 1 Protein 64 Ribonucleic acid 5 Deoxyribonucleic acid 8 Swine [1.203] Phospholipid 18.2 10 Other lipids6 3.0 Protein 74.8 Ribonucleic acid 2.8 Deoxyribonucleic acid 1.2 Erythrocytes, hen [1.20] Phospholipid 13.0 24 Cholesterol 2.5 Protein 75.4 Ribonucleic acid 4.0 Deoxyribonucleic acid 3.8 Thymus, calf [1.19] Phospholipid 16.1 18 Cholesterol 2.2 Protein 70.0 Ribonucleic acid 4.2 Deoxyribonucleic acid 7.5 1Nuclear envelope is localized at two positions after isopyknotic density gradient centrifugation. Major fraction represents approximately 90% of total membrane isolated from gradient. 2Major fraction. 3Mannose is predominant neutral sugar; only trace quantities of galactose detected. Fucose not detected by gas chromatographic analysis of the trimethylsilyl derivatives of envelope carbohydrates. [ref. 15] 4N-Acetyglucosamine, determined as glucosamine after acid hydrolysis, accounts for approximately 95% of total amino sugars. 5Minor fraction. 6Includes cholesterol, triglycerides, and free fatty acids. Contributor: Kasper, Charles B. Valuesare given in percent by weight of all components measured. Data in brackets refer to the column heading in brackets.

References 4. Berezney, R., et al. 1972. J. Biol. Chem. 247:5549–5561. 5. Bornens, M. 1968. C. R. Acad. Sci. 266(D):270–272. 1. Agutter, P. S. 1972. Biochim. Biophys. Acta 255:397– 6. Feldherr, C. M. 1972. Adv. Cell Mol. Biol. 2:273– 401. 307. 2. Berezney, R. 1974. Methods Cell Biol. 8:205–228. 7. Franke, W. W. 1974. Int. Rev. Cytol., suppl. 4:71–236. 3. Berezney, R., et al. 1970. Biochim. Biophys. Acta 8. Franke, W. W. 1974. Phil. Trans. R. Soc. London 203:531–546. B268:67–93.

38 9. Franke, W. W., and U. Scheer. 1974. In H. Busch., ed. 16. Kay, R. R., and I. R. Johnston. 1973. Sub-Cell. Bio- The Cell Nucleus. Academic Press, New York. v. 1: chem. 2:127–167. pp. 219–347. 17. Kay, R. R., et al. 1972. Eur. J. Biochern. 30:145–154. 10. Franke, W. W., et al. 1970. J. Cell Biol. 46:379–395. 18. Matsuura, T., and K. Ueda. 1972. Arch. Biochem. Bio- 11. Jarasch, E.-D., and W. W. Franke. 1974. J. Biol. Chem. phys. 150:440–450. 249:7245–7254. 19. Monneron, A., et al. 1972. J. Cell Biol. 55:104–125. 12. Kashnig, D. M., and C. B. Kasper. 1969. Ibid. 20. Moore, R. J., and J. D. Wilson. 1972. J. Biol. Chem. 244:3786–3792. 247:958–967. 13. Kasper, C. B. 1974. In H. Busch. ed. The Cell Nucleus. 21. Stevens, B. J., and J. Andre. 1969. In A. Lima-de-Faria. Academic Press, New York. v. 1: pp. 349–384. ed. Handbook of Molecular Cytology. North Holland, 14. Kasper, C. B. 1974. Methods Enzymol. 31(A):279– Amsterdam. pp. 837–871. 292. 22. Zbarsky, I. B. 1972. Methods Cell Physiol. 5:167–198. 15. Kasper, C. B. Unpublished. Univ. Wisconsin, McArdle 23. Zbarsky, I. B., et al. 1969. Nature (London) 221:257–259. Laboratory for Cancer Research, Madison, 1975. 24. Zentgraf, H., et al. 1971. J. Biol. Chem. 246:2986–2995.

39 Table 7. Part II. Phospholipid Composition Cell Source Phospholipid Value Reference Liver, bovine Cardiolipin 2.1 2 Lysophosphatidyl choline 1.6 Phosphatidyl choline 54.5 Phosphatidyl ethanolamine 22.5 Phosphatidyl inositol 7.6 Phosphatidyl serine 6.0 Sphingomyelin 5.8 Rat Lysophosphatidyl choline 4.7 6 Phosphatidic acid 5.4 Phosphatidyl choline 58.7 Phosphatidyl ethanolamine 15.0 Phosphatidyl inositol 6.6 Phosphatidyl serine 3.6 Sphingomyelin 6.0 Cardiolipin þ phosphatidic acid 1.5 3 Lysophosphatidyl choline 2.3 Phosphatidyl choline 59.7 Phosphatidyl ethanolamine 18.8 Phosphatidyl inositol þ phosphatidyl serine 14.6 Sphingomyelin 2.7 Swine Lysophosphatidyl choline 1.0 4 Phosphatidic acid 1.0 Phosphatidyl choline 58.2 Phosphatidyl ethanolamine 25.0 Phosphatidyl inositol 8.8 Phosphatidyl serine 4.8 Sphingomyelin 2.0 Erythrocytes, hen Phosphatidyl choline 53.0 5 Phosphatidyl ethanolamine 26.0 Phosphatidyl inositol 7.1 Sphingomyelin 13.0 Thymus, calf Cardiolipin 1.2 1 Lysophosphatidyl choline 1.7 Lysophosphatidyl ethanolamine 4.8 Phosphatidyl choline 68.0 Phosphatidyl ethanolamine 15.0 Phosphatidyl inositol þ phosphatidyl serine 8.0 Sphingomyelin 1.1 Values are given in percent of total phospholipid. Contributor: Kasper, Charles B.

References 3. Khandwala, A. S., and C. B. Kasper. 1971. J. Biol. Chem. 246:6242–6246. 1. Jarasch, E.-D., et al. 1973. Hoppe Seylers Z. Physiol. 4. Kleinig, H. 1970. J. Cell Biol. 46:396–402. Chem. 354:974–986. 5. Kleinig, H., et al. 1971. J. Biol. Chem. 246:2996–3000. 2. Keenan, T. W., et al. 1970. Biochim. Biophys. Acta 6. Lemarchal, P., and M. Bornens. 1969. Bull. Soc. Chim. 203:547–554. Biol. 51:1021–1046.

40 Table 7. Part III. Enzymic Composition Enzyme (Synonym) [Enzyme Cell Source Commission Number] Experimental Conditions Value Reference Liver, bovine NADPH-cytochrome reductase (TPNH-cytochrome c Untreated 0.005 mmole cytochrome c reduced min21 mg protein21 1 reductase) [1.6.2.4] NADH dehydrogenase (DPNH-cytochrome c reductase) Untreated 0.84 mmole cytochrome c reduced min21 mg protein21 1 [1.6.99.3] 21 21 NADH oxidase (DPNH-oxidase) 0.251 mmole O2 consumed min mg protein 1 1 21 21 Cytochrome c oxidase (Cytochrome oxidase) [1.9.3.1] 0.254 mmole O2 consumed min mg protein 1 Cytochromes aþa3 0.045 nmole/mg protein 1 Cytochrome b5 Untreated 0.332 nmole/mg protein 1 Cytochrome P-450 Untreated 0 nmole/mg protein 1 21 21 Glucose-6-phosphatase [3.1.3.9] 1.8 mmole Pi released 15 min mg protein 1 2þ 21 21 Adenosinetriphosphatase (Mg adenosine triphosphatase) 1.7 mmole Pi released 15 min mg protein 1 [3.6.1.3] Rat NADPH-cytochrome reductase (TPNH-cytochrome Untreated 0.0062; 0.0083 mmole cytochrome c reduced min21 15 c reductase) [1.6.2.4] protein21 0.018 mmole cytochrome c reduced min21 protein21 2 0.051 mmole cytochrome c reduced min21 protein21 12 0.104 mmole cytochrome c reduced min21 protein21 7 Phenobarbital treated 0.092 mmole cytochrome c reduced min21 protein21 7 3-Methylcholanthrene treated 0.120 mmole cytochrome c reduced min21 protein21 8 NADH dehydrogenase (DPNH-cytochrome c reductase) Untreated 0.0163; 0.0602mmole cytochrome c reduced min21 mg 15 [1.6.99.3] protein21 0.100 mmole cytochrome c reduced min21 mg protein21 2 0.267 mmole cytochrome c reduced min21 mg protein21 12 0.552 mmole cytochrome c reduced min21 mg protein21 7 Phenobarbital treated 0.401 mmole cytochrome c reduced min21 mg protein21 7 3-Methylcholanthrene treated 0.436 mmole cytochrome c reduced min21 mg protein21 8 21 21 NADH oxidase (DPNH-oxidase) 0.059 mmole O2 consumed min mg protein 3 1 2 3 21 21 Cytochrome c oxidase (Cytochrome oxidase) [1.9.3.1] 0.01 ; 0.08 mmole O2 consumed min mg protein 15 21 21 0.133 mmole O2 consumed min mg protein 3 Cytochromes a þ a3 0 nmole/mg protein 10 0.034 nmole/mg protein 3 Cytochrome b5 Untreated 0.034 nmole/mg protein 2 0.183 nmole/mg protein 7 Phenobarbital treated 0.210 nmole/mg protein 7 3-Methylcholanthrene treated 0.174 nmole/mg protein 8 41 Continued 42 Table 7. Continued Cell Source Enzyme (Synonym) [Enzyme Experimental Conditions Value Reference Commission Number] Cytochrome P-450 Untreated 0.025 nmole/mg protein 2 0.094 nmole/mg protein 3 0.16 nmole/mg protein 8, 9 Phenobarbital treated Fn4 8 Benzopyrene 3-monooxygenase (Aryl hydroxylase) Untreated 259 pmoles hydroxylated benzo[a]pyrene formed 30 min21 7 [1.14.14.2] protein21 328 pmoles hydroxylated benzo[a]pyrene formed 30 min21 11 mg protein21 Phenobarbital treated 208 pmoles hydroxylated benzo[a]pyrene formed 30 min21 7 mg protein21 211 pmoles hydroxylated benzo[a]pyrene formed 30 min21 11 mg protein21 3-Methylcholanthrene 5428 pmoles hydroxylated benzo[a]pyrene formed 11 treated 30 min21 mg protein21 21 212 Glucose-6-phosphatase [3.1.3.9] 0.042 mmole Pi released 15 min mg protein 15 21 21 1.35 mmoles Pi released 15 min mg protein 5 21 21 2.4 mmoles Pi released 15 min mg protein 12 5 2 21 21 3.82 ; 5.32 mmoles Pi released 15 min mg protein 6 21 21 Mannose-6-phosphatase 1.12 mmoles Pi released 15 min mg protein 5 21 21 Inorganic pyrophosphatase [3.6.1.1] 1.12 mmoles Pi released 15 min mg protein 5 2þ 5 2 21 21 Adenosinetriphosphatase (Mg adenosine triphosphatase) 0.433 ; 0.591 mmole Pi released 15 min mg protein 6 21 21 [3.6.1.3] 1.4 mmoles Pi released 15 min mg protein 2 2 3 21 21 3.5 ; 5.9 mmoles Pi released 15 min mg protein 15 3-Methyl-4-monomethylamino-azobenzene N-demethylase Untreated 2.6 nmoles 3-methyl-4-aminoazobenzene formed 30 min21 7 mg protein21 Phenobarbital treated 3.1 nmoles 3-methyl-4-aminoazobenzene formed 30 min21 7 mg protein21

Swine NADPH-cytochrome reductase (TPNH-cytochrome c Untreated 0.021 mmole cytochrome c reduced min21 mg protein21 2 reductase) [1.6.2.4] NADH dehydrogenase (DPNH-cytochrome c reductase) Untreated 0.045 mmole cytochrome c reduced min21 mg protein21 2 [1.6.99.3] Cytochrome bs Untreated 0.026 nmole/mg protein 2 Cytochrome P-450 Untreated 0.032 nmole/mg protein 2 21 21 Glucose-6-phosphatase [3.1.3.9] 0.025 mmole Pi released 15 min mg protein 2 2þ 21 21 Adenosinetriphosphatase (Mg adenosine triphosphatase) 1.7 mmoles Pi released 15 min mg protein 2 [3.6.1.3] Continued Table 7. Continued Cell Source Enzyme (Synonym) [Enzyme Experimental Conditions Value Reference Commission Number]

Erythrocytes, hen NADPH-cytochrome reductase (TPNH-cytochrome c Untreated . 0 mmole cytochrome c reduced min21 mg protein21 16 reductase) [1.6.2.4] NADH dehydrogenase (DPNH-cytochrome c reductase) Untreated 0.17 mmole cytochrome c reduced min21 mg protein21 16 [1.6.99.3] 1 21 21 Cytochrome c oxidase (Cytochrome oxidase) [1.9.3.1] 0 mmole O2 consumed min mg protein 16 21 21 Glucose-6-phosphatase [3.1.3.9] 0 mmole Pi released 15 min mg protein 16 2þ 21 21 Adenosinetriphosphatase (Mg adenosine triphosphatase) 0.025 mmole Pi released 15 min mg protein 16 [3.6.1.3]

6 2 Thymus Calf Cytochromes a þ a3 0.021 ; 0.058 nmoles/mg protein 13 Rat NADPH-cytochrome reductase (TPNH-cytochrome c Untreated 0 mmole cytochrome c reduced min21 mg protein21 4 reductase) [1.6.2.4] NADH dehydrogenase (DPNH-cytochrome c reductase) Untreated 0.017 mmole cytochrome c reduced min21 mg protein21 4 [1.6.99.3] 21 21 NADH oxidase (DPNH-oxidase) 0.014 mmole O2 consumed min mg protein 3 1 21 21 Cytochrome c oxidase (Cytochrome oxidase) [1.9.3.1] 0.037 mmole O2 consumed min mg protein 3 Cytochromes a þ a3 0.020 nmole/mg protein 3 21 21 Glucose-6-phosphatase [3.1.3.9] 0.015 mmole Pi released 15 min mg protein 3 2þ 21 21 Adenosinetriphosphatase (Mg adenosine triphosphatase) 0.78 mmole Pi released 15 min mg protein 3 [3.6.1.3]

Ventral prostate, rat TPNH-D4-3-ketosteroid 5a-reductase 224 pmoles stanolone7 formed hr21 mg protein21 14 1For a discussion of the likelihood that cytochrome c oxidase is not an intrinsic nuclear envelope enzyme, consult reference 3. 2Membrane density = 1.19 g/ml. 3Membrane density = 1.16 g/ml. 4Level of cytochrome P-450 is not increased upon administration of phenobarbital. 5Membrane density = 1.17 g/ml. 6Membrane density = 1.21 g/ml. 7Synonym: Dihydrotestosterone. Value: Pi ¼ inorganic orthophosphate. Data in brackets refer to the column heading in brackets. Contributor: Kasper, Charles B. 43 References 9. Kasper, C. B. 1974. Methods Enzymol. 31(A):279– 292. 1. Berezney, R., et al. 1972. J. Biol. Chem. 247: 10. Khandwala, A. S., and C. B. Kasper. 1972. Fed. Proc. 5549–5561. 31:873 (Abstr. 3735). 2. Franke, W. W., et al. 1970. J. Cell Biol. 46: 379–395. 11. Khandwala, A. S., and C. B. Kasper. 1973. Biochem. 3. Jarasch, E-D., and W. W. Franke. 1974. J. Biol. Chem. Biophys. Res. Commun. 54:1241–1246. 249:7245–7254. 12. Kay, R. R., et al. 1972. Eur. J. Biochem. 30:145–154. 4. Jarasch, E-D., et al. 1973. Hoppe Seylers Z. Physiol. 13. Matsuura, T., and K. Ueda. 1972. Arch. Biochem. Chem. 354:974–986. Biophys. 150:440–450. 5. Kartenbeck, J., et al. 1973. Exp. Cell Res. 81:175–194. 14. Moore, R. J., and J. D. Wilson. 1972. J. Biol. Chem. 6. Kashnig, D. M., and C. B. Kasper. 1969. J. Biol. Chem. 247:958–967. 244:3786–3792. 15. Zbarsky, I. B., et al. 1969. Nature (London) 221:257– 7. Kasper, C. B. 1971. Ibid. 246:577–581. 259. 8. Kasper, C. B. 1974. In H. Busch. ed. The Cell Nucleus. 16. Zentgraf, H., et al. 1971. J. Biol. Chem. 246:2986– Academic Press, New York. v. 1, pp. 349–384. 2995.

Tables adapted by kind permission of Federation of American Societies for Experimental Biology (FASEB) from Biological Handbooks. I. Cell Biology (Altman P.L. and Katz DD; 1976), Bethesda, Maryland.

44