DELAYED FORMATION OF CHROMOSOME ABERRATIONS IN MOUSE PACHYTENE SPERMATOCYTES TREATED WITH TRIETHYLENEMELAMINE (TEM)
|
|
- Margaret Hudson
- 5 years ago
- Views:
Transcription
1 DELAYED FORMATION OF CHROMOSOME ABERRATIONS IN MOUSE PACHYTENE SPERMATOCYTES TREATED WITH TRIETHYLENEMELAMINE (TEM) W. M. GENEROSO,* M. KRISHNA,+,2 R. E. SOTOMAYOR+ AND N. L. A. CACHEIRO* +Biology Division, Oak Ridge National Laboratory and +University of Tennessee, Oak Ridge Graduate School of Biomedical Sciences, Oak Ridge, Tennessee Manuscript received July 21, 1976 Revised copy received October 6, 1976 ABSTRACT Induction of chromosome aberrations in pachytene spermatocytes of mice by 2 mg/kg TEM was compared with induction by 400 R X rays. These doses induced comparably high dominant lethal effects in pachytene spermatocytes of mice. Cytological analysis at diakinesis-metaphase 1 stage showed that whereas 76.4% of the cells treated with X rays at pachytene stage had aberrations, the frequencies observed in two TEM experiments were only 0.8 and 2.2%. On the other hand, 5% of the progeny from TEM-treated pachytene spermatocytes were found to be translocation heterozygotes. This is the first report on the recovery of heritable translocations from treated spermatocytes of mice. The aberration frequencies observed for TEM in diakinesis-metaphase I were much too low to account for all the lethal mutations and heritable translocations. Thus, the formation of the bulk of aberrations induced by TEM in pachytene spermatocytes was delayed--a marked contrast to the more immediate formation of X-ray-induced aberrations. It is postulated that the formation of the bulk of TEM-induced aberrations in pachytene spermatocytes and in certain postmeiotic stages occurs sometime during spermiogenesis, and not through the operation of postfertilization pronuclear DNA synthesis. UALITATIVELY, there is generally no difference in variolus germ-cell Q stages of mice between measurable chromosome breakage effects of ionizing radiations and of the many mutagenic alkylating chemicals. Dominant lethal mutations, heritable translocations, heritable inversions, sex-chromosome loss, and various types of cytologically detectable aberrations are end points that are common for both ionizing radiations and clastogenic alkylating agents. Questions then arise about the nature of events that lead to the formation of chemically induced chromosomal aberrations in mouse germ cells, and about the similarity of mechanisms involved to those for radiation. Both X rays and TEM induce dominant lethal mutations in pachytene spermatocytes of mice. There is strong evidence that dominant lethal mutations induced Research jointly sponsored by the National Center for Toxicological Research and de U. S. Energy Research and and Development Administration under contract with Union Carbide Corporation. a Postdoctoral investigator supported by Subcontract No from de Biology Division of Oak Ridge National Laboratory to the University of Tenliessee. Genetics 85: January, 1977
2 66 w. M. GENEROSO et al. in this germ-cell stage by X rays arise from induced chromosome breakage. OAKBERG and DIMINNO (1960) found an increased number of anaphase I and anaphase I1 cells with bridges, and WENNSTROM (1971), MCGAUGHEY and CHANG (1973), and TSUCHIDA and UCHIDA (1975) found increased frequency of structural aberrations in diakinesis-metaphase I cells following gamma- or X-ray treatment of primary spermatocytes. No such information was available for TEM. Nor was there information on the inducibility by any mutagen of heritable reciprocal translocations in meiotic spermatocytes of mice. Accordingly, the present study was conducted to determine the nature of TEM-induced dominact lethal mutations in pachytene spermatocytes by cytological examination of diakinesis-metaphase I cells and by scoring for transmissible translocations. A comparative study of the effects of TEM and X rays revealed contrasting mechanisms between the two agents in the formation of chromosome aberrations. MATERIALS AND METHODS (101 x C3H)F, hybrid male mice, approximately 12 weeks old, were treated intraperitoneally with 2mg/kg of TEM or exposed to 400 R of acute X rays (partial body irradiation). Control mice were given comparable volumes of Hanks balanced salt solution. In the dominant lethal study, analysis was done on 10- to 12-week-old (C3H x C57BL)F1 females mated on days 25 to 28 after treatment of males. In the heritable translocation study, (SEC x C57BL)F, females mated to TEM-treated males during this posttreatment period were allowed to go to term. Female progeny were discarded, while the males were tested for translocation heterozygosity at maturity (see GENEROSO et al for procedure). Two groups of mice were killed for analysis of diakinesis-metaphase I spermatocytes. The first group was composed of only TEM-treated males, while the second group included TEMand X-ray-treated and control males. Two males per treatment were killed daily at intervals 3 to 7 days postreatment. The two testes of each male were prepared separately, using the air-dry technique of EVANS, BRECKON and FORD (1964.). Twenty-five cells were scored per testis. It is assumed that TEM had no effect on the progression of spermatogenesis and that the cells analyzed at diakinesis-metaphase I stage are the same as those studied in the dominant lethal and heritable translocation experiments at the time of treatment. RESULTS Data presented in Table 1 show that the doses of TEM and X rays used generally cause comparable reductions in pregnancy rate and number of living embryos. That dominact lethal mutations were induced by the two agents is clearly demonstrated by the high frequencies of dead implantations. In the case of X rays, it is known that killing of spermatocytes, in addition to dominant TABLE 1 Comparable dominant lethal effects of X ray and TEM in pachyiene spermatocytes Number of Pregnancy Number of Number of living Dead Tie ilnieut female; mated rate (%) Implants (avg) embryos (avg) implants (%I Control TEM, 2 mg/kg X ray, 400 R
3 HERITABLE TRANSLOCATIONS IN SPERMATOCYTES 67 lethality, contributes to the reduction in litter size (OAKBERG and DIMINNO 1960). Similarly, the reductions in number of living embryos and pregnancy rate after treatment with 2 mg/kg of TEM are partially attributable to killing of pachytene spermatocytes. The rate of fertilization (as judged by the presence of at least one pronucleus) in females mated 25 to 28 days after "EM treatment of males was 42%, compared with 84% for controls (total eggs examined were 188 and 103, respectively). Of the 247 male progeny tested for heritable translocations, 13 (5%) were translocation heterozygotes. The rate of 5% is significantly higher (P < 0.01) than the spontaneous rate of 0.091% (4 translocations in 4392 mice tested) pooled from previous experiments (GENEROSO et al., in press). Of the 13 translocation heterozygotes, all confirmed cytologically, IO were partially sterile and 3 were completely sterile. Results of the cytological analysis of diakinesis-metaphase I spermatocytes are shown in Table 2. It is clear from these data that although a very high frequency TABLE 2 Cytological analysis of diakinesis-metaphase I spermatocytes Posttreatment Number of Number of cells Number of cells Cells with Treatment interval (days) cells scored with fragments with exchanges aberrations (%) Control _ X ray, 400 R TEM, 2 mg/kg ' TEM, 2 mg/kg* ' * Replicate of the above TEM study.
4 68 w. M. GENEROSO et al. of cells treated with X rays at pachytene stage had fragments and/or interchanges, only a small increase in aberration was found with TEM. DISCUSSION The results show that TEM and X rays are alike in the sense that chromosome breakage resulted from treatment of pachytene spermatocytes. In the case of X rays, chromosome breakage was manifested by dominant lethal mutations and by the presence of chromosome aberrations in diakinesis-metaphase I spermatocytes. In the case of TEM, dominant lethal mutations were also clearly induced, but analysis of diakinesis-metaphase I spermatocytes did not give proof that chromosome breakage was the underlying cause. The frequency of observed abcrrations was much too low to account for all the lethals. The clear-cut proof that breakage was induced came rom data on heritable translocations. The very low frequency of aberrations in diakinesis-metaphase I stage and the recovery of heritable translocations in the "EM study provide the first evidence, for any chemical mutagen, that the formation of the bulk of breaks in pachytene spermatocytes of mice is delayed, a marked contrast to the more immediate formation of X-ray-induced breakage. Such a difference between X rays and a chemical mutagen is also evident in mouse dictyate oocytes. SEARLE and BEECHEY ( 1974) observed a high incidence of aberrations when X-irradiated dictyate oocytes were analyzed in metaphase I. BREWEN and PAYNE (in press) on the other hand, using methyl methanesulfonate (MMS), did not find a significant increase over controls in the number of aberrations found in metaphase I. However, they found a marked increase in aberrations when first-cleavage metaphase cells were scored, which indicates that formation of breakage in MMS-treated dictyate oocytes is also delayed, as ir! the present results with pachytene spermatocytes treated with TEM. Similar persistence of TEM-induced premutational lesions in the mouse postmeiotic male germ cells was suggested to be the reason for the premature condensation of certain regions of chromosomes and for the presence of chromatid-type aberrations in the second and third cleavage divisions (MATTER and JAEGER 1975). However, direct evidence for this reasoning is lacking. The present study. indeed, shows that a reasonably high frequency of reciprocal translocations was recovered after treatment of meiotic spermatocytes with a chemical mutagen. This finding is significant in view of the observation from X-ray studies on mouse spermatogonia stem cells that strongly indicates that, for a hitherto unknown reason, the transmission of balanced exchanges is markedly lower than that expected on the basis of the frequency of multivalent association at diakinesjs-metaphase I (BREWEN, PRESTON and GENEROSO 1974; FORD et al. 1969; GENEROSO, CAIN and HUFF 1974). Because of this and the fact that the interchanges induced by X rays in pachytene spermatocytes are of the chromatid type, it is expected that, relative to dominant lethals, the frequency at which reciprocal translocations are recovered among progeny from treated pachytene spermatocytes is lower for X rays than for TEM. On the ather hand,
5 HERITABLE TRANSLOCATIONS IN SPERMATOCYTES 69 it seems likely that the reason heritable translocations were recovered at a frequemy of 5% after TEM treatment of pachytene spermatocytes was that the formation of aberrations was delayed-thus the mechanism may be similar to that of TEM in certain postmeiotic stages. We are presently studying the transmission of X-ray-induced translocations in pachytene spermatocytes. The question which brings into focus the mechanism of aberration formation is, at what point after TEM treatment of pachytene spermatocytes does formation of aberrations actually occur? It is reasonable to assume that the formation of aberrations is initiated by the TEM alkylation of DNA. Following this reaction, does the actual aberration formation occur as a consequence of normal DNA synthesis, as originally suggested by EVANS and SCOTT (1964) for maleic hydrazide effects in Vicia faba, or does it occur independently of DNA synthesis, through the action of various enzyme mechanisms? With respect to the latter possibility, two points need to be stated. First, an intervening round of DNA synthesis does not appear to be necessarily a prerequisite for the formation of exchanges. This is borne out by the high frequencies of exchange aberrations found in diakinesis-metaphase I spermatocytes following X-ray treatment of pachytene spermatocytes (Table 2) and the increase in the number of chromatid exchanges at the first mitosis of 5-bromodeoxyuridine-treated and G,-illuminated cells that were fixed at early periods (when there were very few S cells). Second, the formation of UV- or 5-bromodeoxyuridine-induced chromatid deletions in G,, X, and isochromatid deletions in GI, XI and G,, X, V,,B Chinese hamster tissue culture cells was postulated to be mediated by a single-strand nuclease and by this enzyme. plus a recombination or postreplication repair mechanisms respectively (GRIGGS and BENDER 1973; BENDER, BEDFORD and MITCHELL 1973; BENDER, GRIGGS and WALKER 1973). In other words, does the conversion into aberrations of the TEM premutational lesion placed in pachytene DNA occur during the postmeitoic stages, or after fertilization but before pronuclear DNA synthesis, or during pronuclear DNA synthesis? More research is needed before w2 can satisfactorily answer this important question, but we can speculate on the possibilities. The possibility that the formation of aberrations occurs sometime during spermiogenesis through the action of various repair enzymes, and not through the operation of postfertilization pronuclear DNA synthesis, is favored by the following arguments: First, it is widely known that the formation of aberrations induced by alkylating chemicals in various somatic cells is dependent upon DNA synthesis, and the aberrations observed at first mitosis are of the chromatid type. If such is the case in male meiotic and certair? postmeiotic stages, the great majority of sterile and partially sterile male progeny. classified as translocation heterozygotes, from male parents treated postmeiotically with either TEM or ethyl methanesulfonate (EMS), are expected to be mosaics. In our previous TEM and EMS studies (GENEROSO et al. 1974; GENEROSO et al., in press), in which male progeny from treated early spermatozoa or midspermatids were tested for translocation heterozygosity, we found no evidence of gonadal mosaicism. In one futile attempt to
6 70 w. M. GENEROSO et al. find gonadal mosaics, spermatogonial metaphases of three TEM-induced translocation heterozygotes were analyzed. That at least the great majority of translocation heterozygotes induced postmeiotically by TEM or EMS were not gonadal mosaics is also indicated by the very close similarity in reproductive performance between these traiislocation heterozygotes and those induced at the same staqes by X rays (GENEROSO et al., in press). About one-third of EMS-, TEM-, or X-ray-induced male translocation heterozygotes were completely sterile, and two-thirds were partially sterile. The average numbers of living embryos of females mated to EMS-, TEM-, and X-ray-induced partially sterile males and killed at inidpregnancy were 44, 43, and 44% that of females mated to normal males. If TEM- or EMS-induced translocation heterozygotes were mosaics, we would have expected relatively fewer complete steriles and relatively higher fertility among partial steriles induced by chemical mutagens than among those induced by X rays, because X-ray-induced translocations in male postmeiotic stages are almost certainly not mosaics. CACHEIRO, RUSSELL and SWART- OUT (1974, personal communication), who have been studying the cytogenetic nature of male sterility. have not found a single case of mosaicism in kidney cells of male sterile translocation heterozygotes induced in male postmeiotic stages by X-ray, TEM, EMS, isopropyl methanesulfonate, or cyclophosphamide. With the expectation that chromatid interchanges will result in gonadal mosaics if, indeed, premutational lesions placed in meiotic and certain postmeiotic stages are converted into aberrations during postfertilization DNA synthesis, one would expect lower frequencies of EMS- or TEM-kduced heritable translocations relative to X rays. Data actually show that in early spermatozoa or midspermatids, rates of 32 and 29% (which is about the highest frequency we found for X rays as well) were induced by EMS and TEM, respectively (GENEROSO et al. 1974; GENEROSO et al., in press). Furthermore, the relative frequencies of dominant lethals and heritable translocations for these stages appear to be the same for X rays as for the two chemicals. Secopd, LANG and ADLER (in press) observed a much higher frequency of transmitted translocations induced by MMS in the postmeiotic stages than would be expected from the frequency of chromatid interchanges observed by BREWEN et al. (1975). The latter investigators found only two cells (less than 1 %) with chromatid interchanges that may result in viable mosaics, while the former found % translocation heterozygotes among 250 progeny tested. Since the dose used by LANG and ADLER produced at most 60% dominant lethals, the frequency of heritable translocations based on all conceptuses is calculated to be at least %. It should be noted that among zygotes with complete symmetrical chromatid interchanges we would expect the balanced combination to appear in only 50% of the two-cell eggs, if we assume random segregation. Thus, the frequency of first cleavage male pronuclear metaphases with complete symmetrical chromatid interchanges should be at least %. That the MMS-induced translocation heterozygotes are not mosaics is clearly demonstrated in LANG and ADLER'S study by the close similarity in the frequency of multivalents at diakinesis-metaphase I between the F, sons and their respec-
7 HERITABLE TRANSLOCATIONS IN SPERMATOCYTES 71 tive sires. There was only one exception in which there was considerable difference between the F, and the F, progeny, but this particular F, sire was not likely to be mosaic because he had a higher frequency of multivalents than his F, son. Thus, it is more likely that the translocation heterozygotes induced by MMS, like those induced by EMS, TEM, and X-ray, came from chromosome and not from chromatid interchanges--i.e., the formation of aberrations was completed prior to pronuclear DNA synthesis. It may be argued that if, indeed, the gonadal tissues arise from a single cell at any point during development (excluding the zygote itself), then no gonadal mosaicism would be expected. On the contrary, there is strong evidence which indicates that both the inner cell mass, from which all fetal cells arise, and the gonad are derived from few cells. At the embryo level, GARDNER (1975) recently reviewed the question of whether differentiation of blastomeres into trophoblast or inner cell mass depends on the segregation of cytoplasmic determinants present in the undivided egg (two distinct cytoplasmic regions in the egg segregate into the trophoblast and inner cell mass during cleavage) or on the interrelations of blastomeres during cleavage. Present data overwhelmingly favor the latter hypothesis. and it is thought that the decisive determinative events leading to differentiation of blastomeres into trophoblast or inner cell mass occur between the 8- and 16-cell stages. At the gonad level, RUSSELL (1964) concluded that there is no such thing as purity of germ line, or even an early separation of cell lineages in mammals. This conclusion was based on her studies with a large class of half-mutant animals (mosaics for mutation at any of five coat-color loci) in which both the germ line and the coat were a mixture of cell types. From the foregoing, the expectation that mosaics will be produced if the formation of aberrations occurs during pronuclear DNA synthesis seems unavoidable. Studies are now under way in our laboratory to obtain direct genetic evidence for the time of formation of aberrations after chemical treatment of various male germ-cell stages and to relate this genetic information to molecular mechanisms of aberration formation. LITERATURE CITED BENDER, M. A., J. S. BEDFORD and J. A. MITCHELL, 1973 Mechanisms of chromosomal aberration production. 11. Aberrations induced by 5-bromodeoxyuridine and visible light. Mutat. Res. 20: BENDER, M. A., H. G. GRIGGS and P. L. WALKER, 1973 Mechanisms of chromosomal aberration production. I. Aberration induction by ultraviolet light. Mutat. Res. 20: BREWEN, J. G. and H. S. PAYNE. Studies on chemically-induced dominant lethality: 11. Cytogenetic studies of MMS-induced dominant lethality in maturing dictyate mouse oocytes. Mutat. Res., in press. BREWEN, J. G., H. S. PAYNE, K. P. JONES and R. J. PRESTON, 1975 Studies on chemicallyinduced dominant lethality: I. The cytogenetic basis of MMS-induced dominant lethality in postmeiotic male germ cells. Mutat. Res. 33: BREWEN, J. G., R. J. PRESTON and W. M. GENEROSO, 1974 X-ray-induced translocations: Comparison between cytologically observed and genetically recovered frequencies. Biol. Div. Annu. Prog. Rep., June 30,1974,ORNL-4993, pp
8 72 w. M. GENEROSO et al. CACHEIRO, N. L. A., L. B. RUSSELL and M. S. SWARTOUT, 1974 Translocations, the predominant cause of total sterility in sons of mice treated with mutagens. Genetics 76: EVANS, E. P., G. BRECKON and C. E. FORD, 1964 An air-drying method for meiotic preparations from mammalian testes. Cytogenetics 3 : EVANS, H. J. and D. SCOTT, 1964 Influence of DNA synthesis on the production of chromatid aberrations by X-rays and maleic hydrazide in Vicia faba. Genetics 49: FORD, C. E., A. G. SEARLE, E. P. EVANS and B. J. WEST, 1969 Differential transmission of translocations induced in spermatogonia of mice by irradiation. Cytogenetics 8: GARDNER, R., 1975 Origins and properties of trophoblast, In: Immunobiology of Trophoblast, pp Edited by R. G. EDWARDS, C. W. S. HOWE and M. A. JOHNSON, Cambridge University Press, Cambridge. GENEROSO, W. M., K. T. CAIN and S. W. HUFF, 1974 Dose effects of acute X-rays on induction of heritable reciprocal translocations in mouse spermatogonia. Biol. Div. Annu. Prog. Rep., June 30,1974,ORNL-4993, pp GENEROSO, W. M., K. T. CAIN, S. W. HUFF and D. G. GOSSLEE. Inducibility by chemical mutagens of heritable translocations in male and female germ cells of mice, In: Advances in Modern Toxicology, Vol. 1. Edited by W. G. FLAMM and M. A. MEHLMAN, Hemisphere Publ. Co., Washington, D. C., in press. GENEROSO, W. M., W. L. RUSSELL, S. W. HUFF, S. K. STOUT and D. G. GOSSLEE, 1974 Effects of dose on the induction of dominant lethal mutations and heritable translocations with ethyl methanesulfonate in male mice. Genetics 77: GRIGGS, H. G. and M. A. BENDER, 1973 Photoreactivation of ultraviolet-induced chromosomal aberrations. Science 179: LANG, R. and I. ADLER. Heritable translocation test and dominant lethal assay in male mice with methyl methanesulfonate. Mutat. Res., in press. MCGAUGHEY, R. W. and M. C. CHANG, 1973 Initial chromosomal lesions induced by X-irradiating primary spermatocytes of mice. Can. J. Genet. Cytol. 15: MATTER, B. E. and I. JAEGER, 1975 Premature chromosome condensation, structural chromosome aberrations, and micronuclei in early mouse embryos after treatment of paternal postmeiotic germ cells with triethylenemelamine: Possible mechanisms for chemically induced dominant lethal mutations. Mutat. Res. 33: OAKBERG, E. F. and R. L. DIMINNO, 1960 X-ray sensitivity of primary spermatocytes of the mouse. Int. J. Radiat. Biol. 2: RUSSELL, L. B., 1964 Genetic and functional mosaicism in the mouse, In: Role of Chromosomes in Development, pp Edited by MICHAE LOCKE, Academic Press, Inc., New York. SEARLE, A. G. and C. V. BEECHEY, 1974 Cytogenetic effects of X rays and fission neutrons in female mice. Mutat. Res. 24: TSUCHIDA, W. S. and I. A. UCHIDA, 1975 Radiation-induced chromosome aberrations in mouse spermatocytes and oocytes. Cytogenet. Cell Genet. 14: 1-8. WENNSTROM, J., 1971 Effect of ionizing radiation on the chromosomes in meiotic and mitotic cells. Commentat. Biol. Soc. Sci. Fenn. 45: Corresponding editor: S. WOLFF
EFFECTS OF DOSE ON THE INDUCTION OF DOMINANT-LETHAL MUTATIONS WITH TRIETHYLENEMELAMINE IN MALE MICE1
EFFECTS OF DOSE ON THE INDUCTION OF DOMINANT-LETHAL MUTATIONS WITH TRIETHYLENEMELAMINE IN MALE MICE1 B. E. MATTER2 AND W. M. GENEROSO Biology Division, Ouk Ridge National Laboratory, Oak Ridge, Tennessee
More informationEFFECTS OF DOSE ON THE INDUCTION OF DOMINANT-LETHAL MUTATIONS AND HERITABLE TRANSLOCATIONS WITH ETHYL METHANESULFONATE IN MALE MICE1
EFFECTS OF DOSE ON THE INDUCTION OF DOMINANT-LETHAL MUTATIONS AND HERITABLE TRANSLOCATIONS WITH ETHYL METHANESULFONATE IN MALE MICE1 W. M. GENEROSO, W. L. RUSSELL, SANDRA W. HUFF, SANDRA K. STOUT AND D.
More informationthe permanent transmissible ones. Thus, it is highly desirable from a practical standpoint to have a good understanding of the
Environmental Health Perspectives Evaluation of Chromosome Aberration Effects of Chemicals on Mouse Germ Cells by W. M. Generoso * Chromosomal aberrations induced-- by chemicals represent a class of genetic
More informationRECIPROCAL TRANSLOCATIONS AND REPRODUCTIVE CAPACITY IN RABBITS FOLLOWING EXTERNAL GAMMA IRRADIATION
Bulgarian Journal of Veterinary Medicine (2005), 8, No 4, 227232 RECIPROCAL TRANSLOCATIONS AND REPRODUCTIVE CAPACITY IN RABBITS FOLLOWING EXTERNAL GAMMA IRRADIATION S. GEORGIEVA 1, TS. YABLANSKI 1, P.
More informationIM males and females (MUKHERJEE 1965; SCHEWE, SUZUKI and ERASMUS
THE BROOD PATTERN OF MITOMYCIN-C-INDUCED TRANSLOCATIONS IN DROSOPHILA MELANOGASTER MALES: THE EFFECT OF TIME P. T. SHUKLA AND C. AUERBACH Institute of Animal Genetics, Edinburgh, Scotland Manuscript received
More informationA FORMULA TO PREDICT THE TRANSMISSION FREQUENCY OF ACENTRIC FRAGMENTS*
A FORMULA TO PREDICT THE TRANSMISSION FREQUENCY OF ACENTRIC FRAGMENTS* A. V. CARRANO Laboratory of Radiobiology, University of California, San Francisco, California 94122 Manuscript received August 15,
More informationStage-dependent changes of chromosomal radiosensitivity in primary oocytes of the Chinese hamster
Cytogenet. Cell Genet. 30: 174-178 (1981) Stage-dependent changes of chromosomal radiosensitivity in primary oocytes of the Chinese hamster K. M ikamo, Y. Kamiguchi, K. Funaki, S.Sugawara, and H.T ateno
More informationLecture -2- Environmental Biotechnology
Lecture -2-1-General Bioassay in pollution Monitoring 1 1 Genotoxicity test At the early testing stages, the genotoxicity assays for predicting potential heritable germ cell damage are the same as used
More informationGametogenesis. To complete this worksheet, select: Module: Continuity Activity: Animations Title: Gametogenesis. Introduction
Gametogenesis To complete this worksheet, select: Module: Continuity Activity: Animations Title: Gametogenesis Introduction 1. a. Define gametogenesis. b. What cells are gametes? c. What are the two cell
More informationBIOH122 Session 26 Gametogenesis. Introduction. 1. a. Define gametogenesis. b. What cells are gametes?
BIOH122 Session 26 Gametogenesis Introduction 1. a. Define gametogenesis. b. What cells are gametes? c. What are the two cell division processes that occur during the cell cycle? d. Define the cell cycle.
More informationChromosomes. Bacterial chromosomes are circular. Most higher organisms have linear chromosomes with a centromere that attaches them to the spindle
1 Chromosomes Bacterial chromosomes are circular Most higher organisms have linear chromosomes with a centromere that attaches them to the spindle Centromere can be in the center (metacentric), off-center
More informationTransgenerational Transmission of Radiation Damage: Genomic Instability and Congenital Malformation 1
J. Radiat. Res., 47: Suppl., B19 B24 (2006) Transgenerational Transmission of Radiation Damage: Genomic Instability and Congenital Malformation 1 Christian STREFFER* Genomic instability/malformation/transgenerational
More informationChapter 2. Mitosis and Meiosis
Chapter 2. Mitosis and Meiosis Chromosome Theory of Heredity What structures within cells correspond to genes? The development of genetics took a major step forward by accepting the notion that the genes
More informationMeiosis & Sexual Reproduction. AP Biology
Meiosis & Sexual Reproduction 2007-2008 Cell division / Asexual reproduction Mitosis produce cells with same information identical daughter cells exact copies clones same amount of DNA same number of chromosomes
More informationAn overall genetic risk assessment for radiological protection purposes
Journal of Medical Genetics, 1980, 17, 15-20 An overall genetic risk assessment for radiological protection purposes PER OFTEDAL AND A G SEARLE From the Institute of General Genetics, University of Oslo;
More informationLecture 17: Human Genetics. I. Types of Genetic Disorders. A. Single gene disorders
Lecture 17: Human Genetics I. Types of Genetic Disorders A. Single gene disorders B. Multifactorial traits 1. Mutant alleles at several loci acting in concert C. Chromosomal abnormalities 1. Physical changes
More informationTo General Embryology Dr: Azza Zaki
Introduction To General Embryology The Human Development is a continuous process that begins when an ovum from a female is fertilized by a sperm from a male. Cell division, growth and differentiation transform
More informationThe Cell Life Cycle. S DNA replication, INTERPHASE. G 2 Protein. G 1 Normal THE CELL CYCLE. Indefinite period. synthesis. of histones.
Mitosis & Meiosis The Cell Life Cycle INTERPHASE G 1 Normal cell functions plus cell growth, duplication of organelles, protein synthesis S DNA replication, synthesis of histones THE CELL CYCLE M G 2 Protein
More informationEFFECT OF EXTERNAL GAMMA IRRADIATION ON RABBIT SPERMATOGENESIS
22 Trakia Journal of Sciences, Vol. 4, No. 1, pp 22-26, 2006 Copyright 2005 Trakia University Available online at: http://www.uni-sz.bg ISSN 1312-1723 Original Contribution EFFECT OF EXTERNAL GAMMA IRRADIATION
More informationThe Chromosomal Basis of Inheritance
Chapter 15 The Chromosomal Basis of Inheritance PowerPoint Lectures for Biology, Seventh Edition Neil Campbell and Jane Reece Lectures by Chris Romero Overview: Locating Genes on Chromosomes A century
More informationGenomic Instability Induced by Ionizing Radiation
Genomic Instability Induced by Ionizing Radiation Christian Streffer Universitätsklinikum Essen, 45122 Essen, Germany INTRODUCTION In contrast to general assumptions it has frequently been shown that DNA
More informationChromosome Abnormalities
Chromosome Abnormalities Chromosomal abnormalities vs. molecular mutations Simply a matter of size Chromosomal abnormalities are big errors Two types of abnormalities 1. Constitutional problem present
More informationChapter 8: Cellular Reproduction
Chapter 8: Cellular Reproduction 1. The Cell Cycle 2. Mitosis 3. Meiosis 2 Types of Cell Division 2n 1n Mitosis: occurs in somatic cells (almost all cells of the body) generates cells identical to original
More informationTEXT Introduction During evolutionary history of organisms, the genomes of organisms are continuously being rearranged and reshaped.
TEXT Introduction During evolutionary history of organisms, the genomes of organisms are continuously being rearranged and reshaped. These rearrangements may change the position of a segment within a chromosome,
More informationThe Chromosomal Basis of Inheritance
LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson Chapter 15 The Chromosomal Basis of Inheritance
More informationCell Divisions. The autosomes represent the whole body. * Male Sex Chromosomes: XY * Female Sex Chromosomes: XX
Cell Divisions Each Cell (including gonads) has 46 chromosomes (23 pairs of chromosomes: 22 pairs of autosomes, 1 pair of sex chromosomes) which are located in the nucleus). The autosomes represent the
More informationA. Incorrect! All the cells have the same set of genes. (D)Because different types of cells have different types of transcriptional factors.
Genetics - Problem Drill 21: Cytogenetics and Chromosomal Mutation No. 1 of 10 1. Why do some cells express one set of genes while other cells express a different set of genes during development? (A) Because
More informationRadiation Research Society is collaborating with JSTOR to digitize, preserve and extend access to Radiation Research.
Persistent Chromosome Aberrations in Irradiated Human Subjects Author(s): M. A. Bender and P. C. Gooch Reviewed work(s): Source: Radiation Research, Vol. 16, No. 1 (Jan., 1962), pp. 44-53 Published by:
More informationThe Chromosomal Basis of Inheritance
LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson Chapter 15 The Chromosomal Basis of Inheritance
More informationThe Chromosomal Basis of Inheritance
Chapter 15 The Chromosomal Basis of Inheritance PowerPoint Lecture Presentations for Biology Eighth Edition Neil Campbell and Jane Reece Lectures by Chris Romero, updated by Erin Barley with contributions
More informationCOMPARATIVE FREQUENCY OF X-RAY INDUCED CROSSOVER- SUPPRESSING ABERRATIONS RECOVERED FROM OOCYTES AND SPERM OF DROSOPHZLA MELANOGASTER
COMPARATIVE FREQUENCY OF X-RAY INDUCED CROSSOVER- SUPPRESSING ABERRATIONS RECOVERED FROM OOCYTES AND SPERM OF DROSOPHZLA MELANOGASTER RUTH E. THOMAS2 AND PAUL A. ROBERTS Biology Division, Oak Ridge National
More informationThe Chromosomal Basis of Inheritance
Chapter 15 The Chromosomal Basis of Inheritance PowerPoint Lecture Presentations for Biology Eighth Edition Neil Campbell and Jane Reece Lectures by Chris Romero, updated by Erin Barley with contributions
More informationINDUCTION OF DOMINANT LETHAL MUTATIONS IN INSECT OOCYTES AND SPERM BY GAMMA RAYS AND AN ALKYLATING AGENT: DOSE-RESPONSE AND JOINT ACTION STUDIES
INDUCTION OF DOMINANT LETHAL MUTATIONS IN INSECT OOCYTES AND SPERM BY GAMMA RAYS AND AN ALKYLATING AGENT: DOSE-RESPONSE AND JOINT ACTION STUDIES LEO E. LACHANCE' AND MAXWELL M. CRYSTAL Entomology Research
More informationThe Cell Cycle CHAPTER 12
The Cell Cycle CHAPTER 12 The Key Roles of Cell Division cell division = reproduction of cells All cells come from pre-exisiting cells Omnis cellula e cellula Unicellular organisms division of 1 cell reproduces
More informationThe Chromosomal Basis of Inheritance
The Chromosomal Basis of Inheritance Factors and Genes Mendel s model of inheritance was based on the idea of factors that were independently assorted and segregated into gametes We now know that these
More informationChromosome Structure & Recombination
Chromosome Structure & Recombination (CHAPTER 8- Brooker Text) April 4 & 9, 2007 BIO 184 Dr. Tom Peavy Genetic variation refers to differences between members of the same species or those of different
More informationGENETICS - NOTES-
GENETICS - NOTES- Warm Up Exercise Using your previous knowledge of genetics, determine what maternal genotype would most likely yield offspring with such characteristics. Use the genotype that you came
More informationThe Chromosomal Basis of Inheritance
LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson Chapter 15 The Chromosomal Basis of Inheritance
More informationA. Incorrect! Cells contain the units of genetic they are not the unit of heredity.
MCAT Biology Problem Drill PS07: Mendelian Genetics Question No. 1 of 10 Question 1. The smallest unit of heredity is. Question #01 (A) Cell (B) Gene (C) Chromosome (D) Allele Cells contain the units of
More informationGeneral Embryology. School of Medicine Department of Anatomy and Histology School of medicine The University of Jordan
General Embryology 2019 School of Medicine Department of Anatomy and Histology School of medicine The University of Jordan https://www.facebook.com/dramjad-shatarat What is embryology? Is the science that
More informationThe Chromosomal Basis of Inheritance
LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson Chapter 15 The Chromosomal Basis of Inheritance
More informationChapter 10 Chromosomes and Cell Reproduction
Chapter 10 Chromosomes and Cell Reproduction Chromosomes Organisms grow by dividing of cells Binary Fission form of asexual reproduction that produces identical offspring (Bacteria) Eukaryotes have two
More informationLecture: Variability. Different types of variability in Biology and Medicine. Cytological essentials of heritable diseases. Plan of the lecture
Lecture: Variability. Different types of variability in Biology and Medicine. Cytological essentials of heritable diseases Plan of the lecture 1. Notion of variability. Different types of variability.
More informationGONADAL MOSAICISM AS A FACTOR IN DETERMINING THE RATIO OF VISIBLE TO LETHAL, MUTATIONS IN DROSOPHILA'
GONADAL MOSAICISM AS A FACTOR IN DETERMINING THE RATIO OF VISIBLE TO LETHAL, MUTATIONS IN DROSOPHILA' LUOLIN S. BROWNING AND EDGAR ALTENBURG Texas Medical Center, Inc., Houston, Texas, and Rice University,
More informationVariations in Chromosome Structure & Function. Ch. 8
Variations in Chromosome Structure & Function Ch. 8 1 INTRODUCTION! Genetic variation refers to differences between members of the same species or those of different species Allelic variations are due
More informationLesson 1. Quiz (short) Cell cycle Chromosomes Mitosis phases
Lesson 1 Quiz (short) Cell cycle Chromosomes Mitosis phases 2 Cell division is needed for Growth (Mitosis) Repair (Mitosis) Reproduction (Meiosis) 3 Mitosis consists of 4 phases (division of the nuclear
More informationThe bases on complementary strands of DNA bond with each other in a specific way A-T and G-C
1 Bio 1101 Lecture 6 Ch. 8: Cellular Basis of Reproduction 2 3 4 5 6 Cellular Basis of Reproduction & Inheritance In order for an organism to replace dead cells or to grow and produce new cells, existing
More informationmouse, which show a combination of unusual properties. The
EFFECT OF X-RAYS ON THE MUTATION OF t-alleles IN THE MOUSE MARY F. LYON M.R.C. Radiobio!ogica! Research Unit, Harwell, Berkshire Received 18.ix.59 1.!NTRODUCTtON THE t-alleles are a long series of recessive
More informationChapter 4 The Chromosome Theory of Inheritance
Chapter 4 The Chromosome Theory of Inheritance 4-1 Sections to study 4.1 Chromosomes: The carriers of genes 4.2 Mitosis: Cell division that preserves chromosome number 4.3 Meiosis: Cell division that halve
More informationChapter 15 Notes 15.1: Mendelian inheritance chromosome theory of inheritance wild type 15.2: Sex-linked genes
Chapter 15 Notes The Chromosomal Basis of Inheritance Mendel s hereditary factors were genes, though this wasn t known at the time Now we know that genes are located on The location of a particular gene
More informationEmbrionic death of F1 (%) Irradiated Females + Irradiated Males
Hereditary Radiation Effects In Offspring Of the Second and Third Generations After Irradiation Of Both Grandparents: Experimental Studies and Hereditary Radiation Effects In Offspring Of the First Generation
More informationMEIOTIC BEHAVIOR OF ASYMMETRIC DYADS IN THE MALE DROSOPHILA1
MEIOTIC BEHAVIOR OF ASYMMETRIC DYADS IN THE MALE DROSOPHILA1 S. ZIMMERING AND E. B. BENDBOW Dluision of Biological and Medical Sciences, Brown Uniuersity, Prouidence, Rhode Island 02922 Manuscript received
More informationCell Division and Inheritance
Cell Division and Inheritance Continuing life relies on reproduction Individual organism replacing dead or damaged cells Species making more of same species Reproduction Cells divide, grow, divide again
More informationCYTOGENETICS Dr. Mary Ann Perle
CYTOGENETICS Dr. Mary Ann Perle I) Mitosis and metaphase chromosomes A) Chromosomes are most fully condensed and clearly distinguishable during mitosis. B) Mitosis (M phase) takes 1 to 2 hrs and is divided
More informationToxicity and Genotoxicity of Pendimethalin in Maize and Onion
Kasetsart J. (Nat. Sci.) 44 : 1010-1015 (2010) Toxicity and Genotoxicity of Pendimethalin in Maize and Onion Nuttapol Promkaew 1, Puangpaka Soontornchainaksaeng 1 *, Sansern Jampatong 2 and Piangchan Rojanavipart
More informationUnit 2: Reproduction and Development. The Cell Cycle
PAGE : 1 The Cell Cycle Cell Cycle: A continuous series of cell growth and division for a cell. All cells go through a cell cycle of some sort. The cell cycle consists of two stages. a. Growth Phase Diagram
More informationWhy do cells reproduce?
Outline Cell Reproduction 1. Overview of Cell Reproduction 2. Cell Reproduction in Prokaryotes 3. Cell Reproduction in Eukaryotes 1. Chromosomes 2. Cell Cycle 3. Mitosis and Cytokinesis Examples of Cell
More informationExample: Distance in M.U. % Crossing Over Why? Double crossovers
Example: Distance in M.U. % Crossing Over 1 5 10 15 50 80 100 107 Why? Double crossovers 232 .. A B. a b. 1. A fully heterozygous gray-bodied (b+), normal winged (vg+) female F 1 fruit fly crossed with
More informationThe bases on complementary strands of DNA bond with each other in a specific way A-T and G-C
1 Bio 1101 Lecture 6 (Guided Notes) Ch. 8: Cellular Basis of Reproduction 2 3 4 5 6 Cellular Basis of Reproduction & Inheritance In order for an organism to replace dead cells or to grow and produce new
More informationCell Cycle and Mitosis
Cell Cycle and Mitosis Name Period A# THE CELL CYCLE The cell cycle, or cell-division cycle, is the series of events that take place in a eukaryotic cell between its formation and the moment it replicates
More informationMeiosis. 4. There are multiple alleles for the ABO blood group. Why are there only two of these alleles normally present in any one individual?
Name: ate: 1. The diagram shown represents a cell that will undergo mitosis. Which diagrams below best illustrate the nuclei of the daughter cells that result from a normal mitotic cell division of the
More informationKEY CONCEPT Cells have distinct phases of growth, reproduction, and normal functions.
5.1 The Cell Cycle KEY CONCEPT Cells have distinct phases of growth, reproduction, and normal functions. Objective: Cells have distinct phases of growth, reproduction and normal functions. APK: Why do
More informationThe form of cell division by which gametes, with half the number of chromosomes, are produced. Chromosomes
& Karyotypes The form of cell division by which gametes, with half the number of chromosomes, are produced. Homologous Chromosomes Pair of chromosomes (maternal and paternal) that are similar in shape,
More informationThe Cell Cycle. Chapter 10
The Cell Cycle Chapter 10 Why Do Cells Divide? Unicellular 1. Reproduction Multicellular 1. Grow 2. Repair 3. Development/reproduction Types of Division Prokaryotic cells Binary fission = asexual reproduction
More informationEffect of chemo- or radiotherapeutic agents on human sperm: the reproductive needs of young male cancer survivors. Ghamartaj-Hosseini
Effect of chemo- or radiotherapeutic agents on human sperm: the reproductive needs of young male cancer survivors Effect of chemo- or radiotherapeutic agents on human sperm: the reproductive needs of young
More informationSexual Reproduction and Meiosis
Sexual Reproduction and Meiosis Meiosis sexual reproduction! Meiosis makes the cells that are responsible for sexual reproduction Sexual Reproduction Producing a new organism by combining chromosomes from
More informationCreating Identical Body Cells
Creating Identical Body Cells 5.A Students will describe the stages of the cell cycle, including DNA replication and mitosis, and the importance of the cell cycle to the growth of organisms 5.D Students
More informationDNA double-strand break repair of parental chromatin in ooplasm and origin of de novo mutations. Peter de Boer
DNA double-strand break repair of parental chromatin in ooplasm and origin of de novo mutations Peter de Boer Department of Obst.& Gynaecology, Div. Reproductive Medicine Radboud University Nijmegen Medical
More informationChapter 8. The Cellular Basis of Reproduction and Inheritance. Lecture by Mary C. Colavito
Chapter 8 The Cellular Basis of Reproduction and Inheritance PowerPoint Lectures for Biology: Concepts & Connections, Sixth Edition Campbell, Reece, Taylor, Simon, and Dickey Copyright 2009 Pearson Education,
More informationcells divide? Growth Development Repair Asexual reproduction Formation of gametes
mitosis and meiosis cells divide? Growth Development Repair Asexual reproduction Formation of gametes How does a cell know when to divide? the cell cycle A repeating process of cell growth and division
More informationGenetics 275 Examination February 10, 2003.
Genetics 275 Examination February 10, 2003. Do all questions in the spaces provided. The value for this examination is twenty marks (20% of the grade for the course). The value for individual questions
More informationCell Division. The Process of Cell Division Section Section 10.2: The Process of Cell Division 12/8/2010
The Process of Cell Division Section 10.2 Biology B Section 10.2: The Process of Cell Division The student will investigate and understand common mechanisms of inheritance and protein synthesis. Key concepts
More informationSpermatogenesis. What is it and what does it look like? How do hormones regulate spermatogenesis?
Spermatogenesis What is it and what does it look like? How do hormones regulate spermatogenesis? FSH, androgens, growth factors Animal Physiology (Hill, Wise, Anderson): Ch. 15 435-438 1 Spermatogenesis:
More informationLawrence Berkeley National Laboratory Lawrence Berkeley National Laboratory
Lawrence Berkeley National Laboratory Lawrence Berkeley National Laboratory Title Disruption of Maternal DNA Repair Increases Sperm-Derived Chromosomal Aberrations Permalink https://escholarship.org/uc/item/3s33w7ng
More informationNOTES- CHAPTER 6 CHROMOSOMES AND CELL REPRODUCTION
NOTES- CHAPTER 6 CHROMOSOMES AND CELL REPRODUCTION Section I Chromosomes Formation of New Cells by Cell Division New cells are formed when old cells divide. 1. Cell division is the same as cell reproduction.
More informationCytological effects induced by Agil herbicide to onion
Volume 16(1), 6-72, 212 JOURNAL of Horticulture, Forestry and Biotechnology www.journal-hfb.usab-tm.ro Cytological effects induced by Agil herbicide to onion Bonciu Elena 1* 1 University of Craiova, Faculty
More informationA. LIMA-DE-FARIA Institute of Genetics, University of Lund, Sweden. Received June 11, 1962
~ ~~~~ GENETIC INTERACTION IN RYE EXPRESSED AT THE CHROMOSOME PHENOTYPE A. LIMA-DE-FARIA Institute of Genetics, University of Lund, Sweden Received June, 96 FOR our understanding of the organization of
More informationChapter 14 Cellular Reproduction
Chapter 14 Cellular Reproduction Biology 3201 Introduction One of the important life functions of living things is their ability to reproduce. Reproduction depends on the cell. Cells reproduce in order
More information6.3 DNA Mutations. SBI4U Ms. Ho-Lau
6.3 DNA Mutations SBI4U Ms. Ho-Lau DNA Mutations Gene expression can be affected by errors that occur during DNA replication. Some errors are repaired, but others can become mutations (changes in the nucleotide
More informationUnit 4: Cell Division Guided Notes
Unit 4: Cell Division Guided Notes 1 Chromosomes are structures that contain material When Eukaryotes are not dividing, DNA and Proteins are in a mass called: When the cell divides, it condenses and becomes
More informationGenetics Review. Alleles. The Punnett Square. Genotype and Phenotype. Codominance. Incomplete Dominance
Genetics Review Alleles These two different versions of gene A create a condition known as heterozygous. Only the dominant allele (A) will be expressed. When both chromosomes have identical copies of the
More informationLECTURE 32 GENETICS OF INVERSIONS. A. Pairing of inversion genotypes:
LECTURE 32 GENETICS OF INVERSIONS A. Pairing of inversion genotypes: 1. Characteristic inversion loops form only in chromosomal heterozygotes of both para- and pericentric inversions. Based on the inversion
More informationAnimal Development. Lecture 3. Germ Cells and Sex
Animal Development Lecture 3 Germ Cells and Sex 1 The ovary of sow. The ovary of mare. The ovary of cow. The ovary of ewe. 2 3 The ovary. A generalized vertebrate ovary. (Wilt and Hake, Ch 2, 2004) 4 The
More informationBiology is the only subject in which multiplication is the same thing as division
Biology is the only subject in which multiplication is the same thing as division 2007-2008 The Cell Cycle: Cell Growth, Cell Division Ch. 10 Where it all began You started as a cell smaller than a period
More informationX-Ray-Induced Specific-Locus Mutation Rates in Newborn and Young Mice
University of Tennessee, Knoxville Trace: Tennessee Research and Creative Exchange Doctoral Dissertations Graduate School 8-1972 X-Ray-Induced Specific-Locus Mutation Rates in Newborn and Young Mice Paul
More informationThe questions below refer to the following terms. Each term may be used once, more than once, or not at all.
The questions below refer to the following terms. Each term may be used once, more than once, or not at all. a) telophase b) anaphase c) prometaphase d) metaphase e) prophase 1) DNA begins to coil and
More informationThe Cellular Basis of Reproduction and Inheritance
Chapter 8 The Cellular Basis of Reproduction and Inheritance PowerPoint Lectures for! Biology: Concepts and Connections, Fifth Edition! Campbell, Reece, Taylor, and Simon Lectures by Chris Romero Objective:
More informationUnit 4 - Reproduction
Living Environment Practice Exam- Parts A and B-1 1. Which cell process occurs only in organisms that reproduce sexually? A) mutation B) replication C) meiosis D) mitosis 2. Which sequence represents the
More informationTHE TETRASOMIC FOR CHROMOSOME 4 IN DROSOPHILA MELANOGASTER
THE TETRASOMIC FOR CHROMOSOME 4 IN DROSOPHILA MELANOGASTER E. H. GRELL Biology Division, Oak Ridge National Laboratory, Oak Ridge, Tennessee Received March U), 1961 DROSOPHILA with one more or less 4th
More informationPRODUCTION OF HERITABLE PARTIAL STERILITY IN THE MOUSE BY METHYL METHANESULPHONATE
Brit. J. Pharmacol. (1964), 23, 521-528. PRODUCTION OF HERITABLE PARTIAL STERILITY IN THE MOUSE BY METHYL METHANESULPHONATE BY H. JACKSON, M. PARTINGTON AND A. L. WALPOLE From the Christie Hospital and
More informationVARIEGATED-TYPE POSITION EFFECTS IN THE MOUSE
VARIEGATED-TYPE POSITION EFFECTS IN THE MOUSE LIANE BRAUCH RUSSELL AND JEAN W. BANGHAM Biology Division, Oak Ridge National Laboratory,' Oak Ridge, Tennessee Received December 1, 1960 NTIL recently, V-type
More informationGENES AND CHROMOSOMES CHAPTER 5
CHAPTER 5 GENES AND CHROMOSOMES For many years the standard authority on the chromosomes was Wilson s The Cell in Development and Inheritance. The second edition of this work was published in 1900; it
More informationEdinburgh Research Explorer
Edinburgh Research Explorer Meiosis and retrotransposon silencing during germ cell development in mice Citation for published version: Oellinger, R, Reichmann, J & Adams, IR 2010, 'Meiosis and retrotransposon
More informationChapter 13 Pre-Test Question 2
Student View Summary View Diagnostics View Print View with Answers Settings per Student Questions part 1: Keimzellen und Befruchtung Due: 12:00pm on Wednesday, December 7, 2011 Note: You will receive no
More informationMULTIPLE CHOICE QUESTIONS
SHORT ANSWER QUESTIONS-Please type your awesome answers on a separate sheet of paper. 1. What is an X-linked inheritance pattern? Use a specific example to explain the role of the father and mother in
More informationCell Division. Chromosome structure. Made of chromatin (mix of DNA and protein) Only visible during cell division
Chromosome structure Made of chromatin (mix of DNA and protein) Only visible during cell division Chromosome structure The DNA in a cell is packed into an elaborate, multilevel system of coiling and folding.
More informationThe Living Environment
Name: Class: 1. Asexual reproduction primarily involves the process of 1) pollination 3) spermatogenesis 2) mitosis 4) ovulation The Living Environment Ms. Fazio Base your answers to questions 4 and 5
More informationLYMHOCYTE CHROMOSOMAL ABERRATION ASSAY IN RADIATION BIODOSIMETRY
LYMHOCYTE CHROMOSOMAL ABERRATION ASSAY IN RADIATION BIODOSIMETRY Dr. Birutė Gricienė 1,2 1 Radiation Protection Centre 2 Vilnius University Introduction Ionising radiation is a well-known mutagenic and
More informationHuman inherited diseases
Human inherited diseases A genetic disorder that is caused by abnormality in an individual's DNA. Abnormalities can range from small mutation in a single gene to the addition or subtraction of a whole
More information