Abstract. Introduction. Materials and methods

Size: px
Start display at page:

Download "Abstract. Introduction. Materials and methods"

Transcription

1 RBMOnline - Vol 10. No Reproductive BioMedicine Online; on web 15 December 2004 Article Cytogenetic analysis of human somatic cell haploidization Dr Vasiliy Galat received his PhD in 1997 from the Moscow Biotechnology Institute. He spent 2 years in the Max-Delbruck Center for Molecular Medicine, Berlin as a post-doctoral fellow then moved to the Reproductive Genetics Institute, Chicago. His research interests include fertilization, early mammalian development and cell differentiation. Dr Vasiliy Galat V Galat, S Ozen, S Rechitsky, A Kuliev 1, Y Verlinsky Reproductive Genetics Institute, 2825 North Halsted Street, Chicago, IL 60657, USA 1 Correspondence: Tel: ; Fax: ; anverkuliev@hotmail.com Abstract Despite recent interest in the derivation of female and male gametes through somatic cell nuclear transfer, there is still insufficient data on chromosomal analysis of these gametes resulting from haploidization, especially involving a human nuclear donor and recipient oocytes. The objective of this study was to investigate the fidelity of chromosomal separation during haploidization of human cumulus cells by in-vitro matured human enucleated MII oocytes. A total of 129 oocytes were tested 4 7, 8 14, or h after nuclear transfer (NT) followed by electro-stimulation, resulting in 71.3% activation efficiency on average. Haploidization was documented by the formation of two separate groups of chromosomes, originating from either polar body/pronucleus (PB/PN), or only 2PN, which were tested by 5-colour FISH, or DNA analysis for copy number of chromosomes 13, 16, 18, 21, 22 and X. Two PN were formed more frequently than PB/PN, irrespective of incubation time. In agreement with recent reports on mouse oocytes, as many as 90.2% of the resulting haploid sets tested showed abnormal chromosome segregation, suggesting unsuitability of the resulting artificial gametes for practical application at the present time. Keywords: chromosomal aneuploidy, haploidization, human oocytes, nuclear transfer, oocyte activation Introduction Somatic nuclei transferred (NT) into the cytoplasm of nonactivated eggs normally undergo premature chromosome condensation (PCC), with the condensed chromosomes organized into spindle-like structures resembling a metaphase plate (Tarkowski and Balakier, 1980). Activation of these reconstructed oocytes results in the formation of two sets of chromosomes, one of which separates with the extrusion of the polar body (PB). This is the phenomenon of haploidization of somatic cell nuclei induced by meiotic cytoplasm, representing a possible tool for artificial gamete production for infertility treatment (Tsai et al., 2000; Kaneko et al., 2001; Lacham- Kaplan et al., 2001; Tesarik et al., 2001; Tesarik, 2002; Palermo et al. 2002a,b; Nagy, 2004; Takeuchi and Palermo, 2004). The study of chromosomal distribution of cumulus cell nuclei haploidized by mouse oocytes has shown a correct haploid number of chromosomes in the range of 6 23% (Tateno et al., 2003a; Chen et al., 2004; Heindrycks et al., 2004). Fluorescence in-situ hybridization (FISH) analysis of haploidization outcome was also performed in a small sample of six human cumulus cell nuclei transferred into enucleated human oocytes, which showed normal chromosomal distribution (Tesarik et al., 2001). The present paper describes the results of a study on fidelity of chromosome separation resulting from haploidization of human cumulus cells by human oocytes, and demonstrates an extremely high frequency of chromosomal errors in the derived gametes. Materials and methods The method used was based on inducing nuclei of somatic cells to skip the S-phase of the cell cycle and undergo haploidization through introduction into enucleated MII oocytes, allowing the formation of artificial gametes from somatic cells. A total of 129 immature oocytes were used, which were donated by IVF patients who signed an informed consent form approved by the IRB of Reproductive Genetics Institute. These oocytes were matured in vitro and enucleated using UV-luminescence to ensure that all analysed 199

2 chromosomes resulting from haploidization belonged to the somatic cell nuclei transferred. Cumulus cells, presumably at G0 cell stage, were used as the nucleus donors. The cumulus masses were released into HTF medium (Specialty Media, Phillipsburg, NJ, USA) with hyaluronidase (100 IU/ml; type I-S; Sigma, St Louis, MO, USA) for approximately 5 min, and nuclei were mechanically isolated and introduced into ooplasts by injection. The reconstructed oocytes were cultured in 30 µl drops of HTF in 60 mm Petri dishes covered with the embryo tested mineral oil (Sigma), equilibrated by mixing 50 ml of the appropriate medium with 150 ml of oil incubated at 37 C in a 5% CO 2 /air atmosphere. The oocytes were activated at different times after NT, to evaluate the impact of prolonged nucleus remodelling in the ooplast cytoplasm on the accuracy of chromosome separation to 2PN. Group 1 was activated 4 7, group 2, 8 14, and group 3, h after NT. Activation was performed using an electrofusion device (XRONOS-1; Reproductive Genetics Institute, Chicago, IL, USA) in a medium consisting of 0.3 mol/l mannitol, 0.1 mmol/l MgSO 4, 0.05 mmol/l CaCl 2, and 0.1% polyvinylpyrrolidone. The outcome of haploidization was evaluated by testing either 2PN or PB/PN formed after activation. PB/PN were removed and fixed, or deposited into tubes with lysis buffer containing Protenase K (Invitrogen, Carsband, CA, USA) for molecular testing. The reconstructed oocytes were fixed for FISH analysis in a cold 3:1 solution of methanol and acetic acid and air-dried, while PB/PN were placed on slides and fixed in drops of a 3:1 solution of methanol and acetic acid. The fidelity chromosome distribution in PB/PN was tested by fivecolour FISH probes (Vysis, Downers Grove, IL, USA) or DNA analysis, using short tandem repeats (STR) markers for chromosomes 13, 16, 18, 21, 22 and X, in a multiplex nested PCR system. To prove the donor origin of the resulting haploid cells DNA profiles of the donor cumulus cells and recipient oocytes were also tested. For each oocyte nucleus donor pairs, different sets of STR markers were used depending on the genotype of each donor. Statistical analysis was performed using a Fisher exact test of differences between proportions. Results As seen from Table 1, 92 (71.3%) of a total of 129 oocytes surviving microsurgery were activated by electric stimulation, resulting in haploidization with the extrusion of PB in 12 (13%), and formation of 1, 2 or multiple PN in 80 (87%) oocytes. Overall, 41 (31.8%) of these oocytes were with 2PN, which is significantly higher than the number of oocytes with PB/PN (12; 9.3%; P < 0.01). However, this difference was less obvious for group 1 (12.1 versus 24.3%, not significant), compared with groups 2 and 3: (6.8 versus 41.4% for group 2; P < 0.01; 3.8 versus 42.4% for group 3; P < 0.01). Although the resulting pronuclei were similar in appearance to those derived from normal fertilization (Figure 1), there were positional differences from the early stages of development. While pronuclei resulting from fertilization migrate from the periphery towards each other and to the centre (Kunkle and Longo, 1975), the haploidization-induced pronuclei remained closely connected despite growing in volume. However, the pronuclei may be separated from each other microsurgically and used for either subsequent analysis or nuclear transfer. Table 2 presents the results of FISH analysis in the resulting haploid sets performed in 41 reconstructed oocytes, only four of which showed a normal distribution of chromosomes; the remaining 37 (90.2%) were abnormal, with one, two or three errors. Group 2 demonstrated a tendency towards higher fidelity of chromosomal separation compared with group 1 (18.2 versus 11.1%), while no euploidy and generally complex errors were observed in group 3. Although the sample is not sufficiently large to demonstrate statistical significance, there was higher fidelity of chromosomal segregation in the haploidization resulting in PB extrusion in groups 1 and 2 compared with the 2PN pattern (28.6 versus 9.0%). Polymerase chain reaction-based testing of the resulting haploid sets, performed in four reconstructed oocytes, showed no DNA residue from the recipient oocytes, confirming the donor genotype of both pronuclei (Figure 2). However, all the resulting pronuclei showed chromosomal errors, involving unequal numbers of chromatids at least for two chromosomes, with the X chromosome being involved in all the errors detected. Discussion These data demonstrate a relatively high activation rate of 71.3% obtained by electrical stimulation, despite the use of in- Table 1. Efficiency of different activation times after incubation following nuclear transfer (NT)(%). Experimental groups Not activated PB/PN PN 2PN >2PN Total NT (time of nuclear remodelling, h) Group 1 (4 7) 25 (33.8) g 9 (12.1) a 14 (18.9) 18 (24.3) d 8 (10.9) 74 Group 2 (8 14) 6 (20.7) h 2 (6.8) b 8 (27.6) 12 (41.4) e 1 (3.5) 29 Group 3 (15 21) 6 (23.0) i 1 (3.8) c 5 (19.3) 11 (42.4) f 3 (11.5) 26 Groups (4 14) 31 (30.0) 11 (10.7) 22 (21.5) 30 (29.1) 9 (8.7) 103 Groups (4 21) 37 (28.7) 12 (9.3) 27 (20.9) 41 (31.8) 12 (9.3) Superscripts indicate statistical significance of comparisons: PB extrusion versus 2PN formation group 1, not significant (a,d); group 2, P < 0.01 (b,e); group 3, P < 0.01 (c,f). PB extrusion rate between groups, not significant (a,b,c). 2PN formation rate between groups, not significant (d,e,f). NT oocyte activation rate between groups, not significant (g,h,i).

3 a b c d Figure 1. In-vitro development of reconstructed oocytes following haploidization. (a c) Reconstructed by somatic cell nuclei injection MII oocytes 10 h after electrostimulation: two haploidization pronuclei position close to each other from the very beginning, despite volume growth. (d) Isolated pronuclei ( 400). Table 2. Chromosomal aneuploidies (%) after haploidization. Experimental groups Normal 1 error 2 errors 3 errors Total NT (time of nuclei haploid remodelling, h) Group 1 (4 7) 2 (11.1) 9 (50.0) 7 (38.9) 0 18 Group 2 (8 14) 2 (18.2) 1 (9.0) 4 (36.4) 4 (36.4) 11 Group 3 (15 21) 0 4 (33.3) 4 (33.3) 4 (33.3) 12 Groups (4 14) 4 (13.8) 10 (34.5) 11 (37.9) 4 (13.8) 29 Groups (4 21) 4 (9.8) 14 (34.1) 15 (36.6) 8 (19.5)

4 202 Figure 2. Capillary electrophoresis: results of cumulus cell haploidization. (a) Normal segregation of chromosomes 21. Each of the pronuclei obtained from cumulus cells injected into the enucleated recipient MII oocyte has one chromosome 21, based on testing for the STR marker D21S290 located on chromosome 21 (PN1 and PN2), evident from the presence of a single allele 225 or 233 for these markers in each pronucleus. The DNA profiles of donor cumulus cell nuclei (donor nucleus), and recipient oocyte (oocyte nucleus) are presented below, demonstrating the evidence that both resulting pronuclei originated from haploidization. (b) Abnormal segregation of X chromosomes. None of the somatic cell alleles for chromosome X marker is present in a resulting pronucleus (PN1), which is in accordance with both of the chromosome X STR marker DXS8061 alleles 160/162 being detected in the corresponding sister pronucleus (PN2). The DNA profiles of donor cumulus cell nuclei (donor nucleus), and recipient oocyte (oocyte nucleus) are presented below, demonstrating the evidence that both of the resulting pronuclei originated from haploidization.

5 vitro matured oocytes, which would be expected to have lower activation and fertilization rates compared with naturally ovulated oocytes. The activation rate varied between the groups depending on the incubation time after NT, which may be extrapolated as the oocyte age, with the lowest activation rate observed in Group 1, stimulated only 4 7 h after NT. The fact that activation ability of oocytes may increase with oocyte ageing and with MII oocyte cytoplasm maturation to a competent state has previously been reported for other species (Ware et al., 1989). The evidence of haploidization, which occurred in 41.1% of activated oocytes in this study, was documented not only by PB extrusion, but also by 2PN-formation, the remaining activation patterns, including 1PN and multiple PN, obviously not representing haploidization (Table 1). The observed morphological appearance of activated oocytes may depend on the method of activation and the time of nuclear remodelling after NT, with relevant changes of spindle structures described in ageing mouse eggs (Webb et al., 1986). Initially, when the spindle moves towards the centre, reshaping metaphase to an anaphase-like structure, 2PN are formed in the activated eggs, but multiple PN are formed with the spindle becoming scattered and dissolved. In contrast to PB/PN haploidization events reported previously (Tesarik et al. 2001), the majority of the present series are represented by 2PN formation, which may be due to the increased time of nuclear remodelling or electrical stimulation used in this study compared with a sperm stimulation protocol. The fact that the PB extrusion rate after NT may differ depending on activation protocol used has also been shown in other species (Motlik et al., 2002). As seen from the DNA fingerprinting results, the resulting PN originated from the donor nuclei (Figure 2), but the majority (90.3%) were chromosomally abnormal. Therefore, although haploidization of somatic cells may be achieved consistently using MII oocyte cytoplasm, the aneuploidy rate is unacceptably high compared with normal meiosis. It may also be noted that haploidization failure in some instances may be associated with the fact that not all cumulus nuclei may be exactly at the G0 stage, as they were taken from cumulus oocyte complexes of IVF patients. As shown in mice, ovulation induction may induce some cumulus granulosa cells to enter a cell cycle (Schuetz, 1996). It will be necessary to improve the fidelity of chromosome distribution, if the resulting gametes are to be of any practical future use. For example, drug-induced arrest at G2 M cell phase was shown to dramatically increase the fidelity of chromosome separation in budding yeast with mutation in spindle checkpoint (Shonn et al., 2000). A reduced aneuploidy rate with incubation for 8 14 h was also observed in group 2 compared with group 1 (18.2 versus 11.1%), but no further improvement was observed with h post-nt incubation in group 3, leading to a contrasting increase in aneuploidy rate. The possible improvement was also observed in the haploidization resulting in PB/PN compared with 2PN (28.6 versus 11.1%), although the data are too preliminary to confirm this. In conclusion, the available data show that despite previous hopes for using nuclear transfer technology to produce female and male gametes through haploidization of somatic cells, the majority of the resulting haploid cells are chromosomally abnormal. Even if somatic cell haploidization could produce haploids with euploid chromosome sets, the probability of imprinting disorders cannot be excluded (Tateno et al., 2003b). Therefore, despite the feasibility of somatic cell haploidization using MII oocyte cytoplasm, clinical use of the resulting artificial gametes cannot be considered at the present time. References Chen SU, Chang CY, Lu CC et al Microtubular spindle dynamics and chromosome complements from somatic cell nuclei haploidization in mature mouse oocytes and developmental potential of the derived embryos. Human Reproduction 19, Heindrycks B, Lierman S, Van der Elst J, Dhont M 2004 Chromosome number and development of artificial mouse oocytes and zygotes. Human Reproduction 19, Kaneko M, Takeuchi T, Veek LL et al Haploidization enhancement to manufacture human oocytes. Human Reproduction 16 (suppl. 1), 4 5. Kunkle M, Longo FJ 1975 Cytological events leading to the cleavage of golden hamster zygotes. Journal of Morphology 146, Lacham-Kaplan O, Daniels R, Trounson A 2001 Fertilization of mouse oocytes using somatic cells as male germ cells. Reproductive BioMedicine Online 2, Motlik J, Alberio R, Zakhartchenko V et al The effect of activation of mammalian oocytes on remodeling of donor nuclei after nuclear transfer. Cloning Stem Cells 4, Nagy ZP 2004 Clinical prospects of nuclear transfer and somatic cell haploidization: haploidization to produce human embryos: a new frontier for micromanipulation. Reproductive BioMedicine Online 8, Palermo GD, Takeuchi T, Rosenwaks Z 2002a Oocyte-induced haploidization. Reproductive BioMedicine Online 4, Palermo GD, Takeuchi T, Rosenwaks Z 2002b Technical approaches to correction of oocyte aneuploidy. Human Reproduction 17, Schuetz AW, Whittingham DG, Snowden R 1996 Alterations in the cell cycle of mouse cumulus granulosa cells during expansion and mucification in vivo and in vitro. Reproductive Fertility Development 8, Shonn MA, McCarroll R, Murray AW 2000 Requirement of the spindle checkpoint for proper chromosome segregation in budding yeast meiosis. Science 289, Takeuchi T, Palermo G 2004 Clinical prospects of nuclear transfer and somatic cell haploidization: implications of cloning technique for reproductive medicine Reproductive BioMedicine Online 8, Tarkowski AK, Balakier H 1980 Nucleo cytoplasmic interactions in cell hybrids between mouse oocytes, blastomeres and somatic cells. Journal of Embryology and Experimental Morphology 55, Tateno H, Akutsu H, Kamiguchi Y et al. 2003a Inability of mature oocytes to create functional haploid genomes from somatic cell nuclei. Fertility and Sterility 79, Tateno H, Latham KE, Yanagimachi R 2003b Reproductive semicloning respecting biparental origin. A biologically unsound principle. Human Reproduction 18, Tesarik J 2002 Reproductive semi-cloning respecting biparental origin: embryos from syngamy between a gamete and a haploidized somatic cell. Human Reproduction 17, Tesarik J, Nagy ZP, Sousa M et al Fertilizable oocytes reconstructed from patient s somatic cell nuclei and donor ooplasts. Reproductive BioMedicine Online 2, Tsai MC, Takeuchi T, Bedford JM et al Alternative sources of gametes: reality or science fiction? Human Reproduction 15,

6 Ware CB, Barnes FL, Maiki-Laurila M, First NL 1989 Age dependence of bovine oocyte activation. Gamete Research 22, Webb M, Howlett SK, Maro B 1986 Parthenogenesis and cytoskeletal organization in ageing mouse eggs. Journal of Embryology and Experimental Morphology 95, Received 19 October 2004; refereed 5 November 2004; accepted 29 November

Articles Somatic cell haploidization: an update

Articles Somatic cell haploidization: an update RBMOnline - Vol 6. No 1. 60 65 Reproductive BioMedicine Online; www.rbmonline.com/article/717 on web 4 November 2002 Articles Somatic cell haploidization: an update Jan Tesarik obtained his MD degree in

More information

A comparison of the effects of estrus cow. nuclear maturation of bovine oocytes

A comparison of the effects of estrus cow. nuclear maturation of bovine oocytes A comparison of the effects of estrus cow serum and fetal calf serum on in vitro nuclear maturation of bovine oocytes J Spiropoulos, SE Long University of Bristol, School of Veterinary Science, Department

More information

Article Pre-embryonic diagnosis for Sandhoff disease

Article Pre-embryonic diagnosis for Sandhoff disease RBMOnline - Vol 12. No 3. 2006 328-333 Reproductive BioMedicine Online; www.rbmonline.com/article/2100 on web 9 January 2006 Article Pre-embryonic diagnosis for Sandhoff disease Dr Anver Kuliev received

More information

SHORT COMMUNICATION CHICAGO, ILLINOIS. Visualization of Chromosomes in Single Human Blastomeres

SHORT COMMUNICATION CHICAGO, ILLINOIS. Visualization of Chromosomes in Single Human Blastomeres Journal of Assisted Reproduction and Genetics, Vol. 16, No. 3, 1999 SHORT COMMUNICATION CHICAGO, ILLINOIS Visualization of Chromosomes in Single Human Blastomeres The present work describes our results

More information

Articles Polar body-based preimplantation diagnosis for X-linked disorders

Articles Polar body-based preimplantation diagnosis for X-linked disorders RBMOnline - Vol 4. No 1. 38 42 Reproductive BioMedicine Online; www.rbmonline.com/article/384 on web 20 November 2001 Articles Polar body-based preimplantation diagnosis for X-linked disorders Dr Yury

More information

How do living things Sexually Reproduce?

How do living things Sexually Reproduce? How do living things Sexually Reproduce? Besides animals, what other things reproduce sexually? Think of a family that has both biological parents and has 2 or more children #1 Consider what the parents

More information

Organisms that reproduce Sexually are made up of two different types of cells.

Organisms that reproduce Sexually are made up of two different types of cells. MEIOSIS Organisms that reproduce Sexually are made up of two different types of cells. 1. Somatic Cells are body cells and contain the normal number of chromosomes.called the Diploid number (the symbol

More information

Article Duplication of the sperm genome by human androgenetic embryo production: towards testing the paternal genome prior to fertilization

Article Duplication of the sperm genome by human androgenetic embryo production: towards testing the paternal genome prior to fertilization RBMOnline - Vol 14. No 4. 2007 504-514 Reproductive BioMedicine Online; www.rbmonline.com/article/2706 on web 20 February 2007 Article Duplication of the sperm genome by human androgenetic embryo production:

More information

The Cell Life Cycle. S DNA replication, INTERPHASE. G 2 Protein. G 1 Normal THE CELL CYCLE. Indefinite period. synthesis. of histones.

The 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 information

Cell cycle co-ordination in embryo cloning by nuclear transfer

Cell cycle co-ordination in embryo cloning by nuclear transfer Reviews of Reproduction (1996) 1, 40 46 Cell cycle co-ordination in embryo cloning by nuclear transfer Keith H. S. Campbell, Pasqualino Loi*, Pedro J. Otaegui and Ian Wilmut Roslin Institute (Edinburgh),

More information

Article Preimplantation diagnosis and HLA typing for haemoglobin disorders

Article Preimplantation diagnosis and HLA typing for haemoglobin disorders RBMOnline - Vol 11. No 3. 2005 362-370 Reproductive BioMedicine Online; www.rbmonline.com/article/1853 on web 20 July 2005 Article Preimplantation diagnosis and HLA typing for haemoglobin disorders Dr

More information

Unit 4: Cell Division Guided Notes

Unit 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 information

Preimplantation genetic diagnosis: polar body and embryo biopsy

Preimplantation genetic diagnosis: polar body and embryo biopsy Human Reproduction, Vol. 15, (Suppl. 4), pp. 69-75, 2000 Preimplantation genetic diagnosis: polar body and embryo biopsy Luca Gianaroli SISMER, Via Mazzini 12, 40138 Bologna, Italy Scientific Director

More information

NOTES- CHAPTER 6 CHROMOSOMES AND CELL REPRODUCTION

NOTES- 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 information

Mitosis: cell division that forms identical daughter cells with the same number of chromosomes as the parent cell (duplicate and divide)

Mitosis: cell division that forms identical daughter cells with the same number of chromosomes as the parent cell (duplicate and divide) Mitosis: cell division that forms identical daughter cells with the same number of chromosomes as the parent cell (duplicate and divide) Meiosis: cell division that forms daughter cells with half the number

More information

Cell Division. The Process of Cell Division Section Section 10.2: The Process of Cell Division 12/8/2010

Cell 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 information

Development of normal mice from metaphase I oocytes fertilized with primary spermatocytes

Development of normal mice from metaphase I oocytes fertilized with primary spermatocytes Proc. Natl. Acad. Sci. USA Vol. 95, pp. 5611 5615, May 1998 Developmental Biology Development of normal mice from metaphase I oocytes fertilized with primary spermatocytes ATSUO OGURA*, OSAMU SUZUKI*,

More information

Cell Division Questions. Mitosis and Meiosis

Cell Division Questions. Mitosis and Meiosis Cell Division Questions Mitosis and Meiosis 1 10 Do not write outside the box 5 Figure 3 shows a pair of chromosomes at the start of meiosis. The letters represent alleles. Figure 3 E E e e F F f f 5 (a)

More information

Chapter 10 Chromosomes and Cell Reproduction

Chapter 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 information

Lesson 1. Quiz (short) Cell cycle Chromosomes Mitosis phases

Lesson 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 information

SUPPORTING ONLINE MATERIAL

SUPPORTING ONLINE MATERIAL SUPPORTING ONLINE MATERIAL SUPPORTING ONLINE TEXT Efficiency of SCNT Alive fetuses at mid-gestation The rate of viable (beating heart) embryos at day 12.5-14.5 dpc was assessed after sacrifice of foster

More information

The Cell Cycle CHAPTER 12

The 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 information

Part II The Cell Cell Division, Chapter 2 Outline of class notes

Part II The Cell Cell Division, Chapter 2 Outline of class notes Part II The Cell Cell Division, Chapter 2 Outline of class notes 1 Cellular Division Overview Types of Cell Division Chromosomal Number The Cell Cycle Mitoses Cancer Cells In Vitro Fertilization Infertility

More information

Mitosis & Meiosis. Practice Questions. Slide 1 / 68. Slide 2 / 68. Slide 3 / Identify two differences between meiosis and mitosis.

Mitosis & Meiosis. Practice Questions. Slide 1 / 68. Slide 2 / 68. Slide 3 / Identify two differences between meiosis and mitosis. New Jersey Center for Teaching and Learning Slide 1 / 68 Progressive Science Initiative This material is made freely available at www.njctl.org and is intended for the non-commercial use of students and

More information

Mitosis & Meiosis. Practice Questions. Slide 1 / 68. Slide 2 / 68. Slide 3 / 68. Slide 4 / 68. Slide 6 / 68. Slide 5 / 68

Mitosis & Meiosis. Practice Questions. Slide 1 / 68. Slide 2 / 68. Slide 3 / 68. Slide 4 / 68. Slide 6 / 68. Slide 5 / 68 Slide 1 / 68 Slide 2 / 68 New Jersey Center for Teaching and Learning Progressive Science Initiative This material is made freely available at www.njctl.org and is intended for the non-commercial use of

More information

Gametogenesis. 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 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 information

Fertilization failures and abnormal fertilization after intracytoplasmic sperm injection

Fertilization failures and abnormal fertilization after intracytoplasmic sperm injection Fertilization failures and abnormal fertilization after intracytoplasmic sperm injection Sean P.Flaherty 1, Dianna Payne and Colin D.Matthews Reproductive Medicine Unit, Department of Obstetrics and Gynaecology,

More information

Zygotes showing a single pronucleus

Zygotes showing a single pronucleus In vitro development and chromosome constitution of embryos derived from monopronucleated zygotes after intracytoplasmic sperm injection Sílvia Mateo, M.Sc., a Monica Parriego, M.Sc., a Montserrat Boada,

More information

Article Poor development of human nuclear transfer embryos using failed fertilized oocytes

Article Poor development of human nuclear transfer embryos using failed fertilized oocytes RBMOnline - Vol 11. No 6. 2005 740 744 Reproductive BioMedicine Online; www.rbmonline.com/article/1888 on web 10 October 2005 Article Poor development of human nuclear transfer embryos using failed fertilized

More information

Meiosis & Sexual Reproduction. AP Biology

Meiosis & 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 information

(i) List these events in the correct order, starting with D.... (1)... (1)... (1)

(i) List these events in the correct order, starting with D.... (1)... (1)... (1) Q1. (a) Boxes A to E show some of the events of the cell cycle. A Chromatids seperate B Nuclear envelopes disappears C Cytoplasm divides D Chromosomes condense and become visible E Chromosomes on the equator

More information

Unit 2: Reproduction and Development. The Cell Cycle

Unit 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 information

BIOH122 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? 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 information

CHAPTER 8 CELL REPRODUCTION

CHAPTER 8 CELL REPRODUCTION CHAPTER 8 CELL REPRODUCTION CHROMOSOME STRUCTURE Structures in the nucleus that carry genetic information Composed of DNA coiled around proteins called histones Consists of 2 identical parts called sister

More information

Biology 4361 Developmental Biology. October 11, Multiple choice (one point each)

Biology 4361 Developmental Biology. October 11, Multiple choice (one point each) Biology 4361 Developmental Biology Exam 1 October 11, 2005 Name: ID#: Multiple choice (one point each) 1. Sertoli cells a. surround spermatocytes b. are the structural components of the seminiferous tubules

More information

Approaches to Preimplantation Diagnosis

Approaches to Preimplantation Diagnosis Approaches to Preimplantation Diagnosis 2 Introduced only in 1990 as an experimental procedure, preimplantation genetic diagnosis (PGD) is now becoming an established clinical option in reproductive medicine

More information

The form of cell division by which gametes, with half the number of chromosomes, are produced. Chromosomes

The 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 information

Polar Body Approach to PGD. Anver KULIEV. Reproductive Genetics Institute

Polar Body Approach to PGD. Anver KULIEV. Reproductive Genetics Institute Polar Body Approach to PGD Anver KULIEV Reproductive Genetics Institute DISCLOSURE othing to disclose 14 History of Polar Body Approach 14 First proposed in World Health Organization s Document Perspectives

More information

Chapter 8: Cellular Reproduction

Chapter 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 information

Abstract. Introduction. RBMOnline - Vol 11. No Reproductive BioMedicine Online; on web 15 June 2005

Abstract. Introduction. RBMOnline - Vol 11. No Reproductive BioMedicine Online;  on web 15 June 2005 RBMOnline - Vol 11. No 2. 2005 226 231 Reproductive BioMedicine Online; www.rbmonline.com/article/1872 on web 15 June 2005 Article Derivation of a human blastocyst after heterologous nuclear transfer to

More information

Biology 3201 Unit 2A Notes

Biology 3201 Unit 2A Notes Biology 3201 Unit 2A Notes Mitosis Somatic cell: refers to a body cell; anything not a germ (sex) cell Cell cycle: a continuous sequence of cell growth and division The cell cycle consists of two main

More information

Article Obtaining metaphase spreads from single blastomeres for PGD of chromosomal rearrangements

Article Obtaining metaphase spreads from single blastomeres for PGD of chromosomal rearrangements RBMOnline - Vol 14. No 4. 2007 498-503 Reproductive BioMedicine Online; www.rbmonline.com/article/2639 on web 6 February 2007 Article Obtaining metaphase spreads from single blastomeres for PGD of chromosomal

More information

Article Kinetic markers of human embryo quality using time-lapse recordings of IVF/ICSI-fertilized oocytes

Article Kinetic markers of human embryo quality using time-lapse recordings of IVF/ICSI-fertilized oocytes RBMOnline - Vol 17 No 3. 2008 385-391 Reproductive BioMedicine Online; www.rbmonline.com/article/3327 on web 30 July 2008 Article Kinetic markers of human embryo quality using time-lapse recordings of

More information

In vitro development of haploid mouse embryos produced by bisection of one-cell fertilized eggs

In vitro development of haploid mouse embryos produced by bisection of one-cell fertilized eggs /. Embryo!, exp. Morph. Vol. 38, pp. 187-202, 1977 187 Printed in Great Britain In vitro development of haploid mouse embryos produced by bisection of one-cell fertilized eggs ByANDRZEJ K.TARKOWSKI 1 From

More information

The 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. 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 information

Abstract. Introduction. RBMOnline - Vol 8. No Reproductive BioMedicine Online; on web 10 December 2003

Abstract. Introduction. RBMOnline - Vol 8. No Reproductive BioMedicine Online;   on web 10 December 2003 RBMOnline - Vol 8. No 2. 224-228 Reproductive BioMedicine Online; www.rbmonline.com/article/1133 on web 10 December 2003 Article Preimplantation genetic diagnosis for early-onset torsion dystonia Dr Svetlana

More information

Cell Division and Inheritance

Cell 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 information

IVF Michigan, Rochester Hills, Michigan, and Reproductive Genetics Institute, Chicago, Illinois

IVF Michigan, Rochester Hills, Michigan, and Reproductive Genetics Institute, Chicago, Illinois FERTILITY AND STERILITY VOL. 80, NO. 4, OCTOBER 2003 Copyright 2003 American Society for Reproductive Medicine Published by Elsevier Inc. Printed on acid-free paper in U.S.A. CASE REPORTS Preimplantation

More information

Chapter 2. Mitosis and Meiosis

Chapter 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 information

Omnis cellula e cellula

Omnis cellula e cellula Chapter 12 The Cell Cycle Omnis cellula e cellula 1855- Rudolf Virchow German scientist all cells arise from a previous cell Every cell from a cell In order for this to be true, cells must have the ability

More information

Chromosomes and Cell Cycle

Chromosomes and Cell Cycle Chromosomes and Cell Cycle Cell Basics There are trillions of cells in your body Cells are microscopic Cells have DNA inside a structure called the nucleus The nucleus is enclosed by a structure called

More information

In vitro maturation of human preovulatory oocytes reconstructed by germinal vesicle transfer

In vitro maturation of human preovulatory oocytes reconstructed by germinal vesicle transfer FERTILITY AND STERILITY VOL. 71, NO. 4, APRIL 1999 Copyright 1999 American Society for Reproductive Medicine Published by Elsevier Science Inc. Printed on acid-free paper in U.S.A. In vitro maturation

More information

Summary. Mouse eggs were fertilized in vitro, in the presence and

Summary. Mouse eggs were fertilized in vitro, in the presence and THE R\l=O^\LEOF CUMULUS CELLS AND THE ZONA PELLUCIDA IN FERTILIZATION OF MOUSE EGGS IN VITRO A. PAVLOK and ANNE McLAREN Czechoslovak Academy of Sciences, Laboratory of Animal Genetics, Libechov, Czechoslovakia,

More information

Case report Successful pregnancy after ICSI with strontium oocyte activation in low rates of fertilization

Case report Successful pregnancy after ICSI with strontium oocyte activation in low rates of fertilization RBMOnline - Vol 13 No 6. 2006 801-806 Reproductive BioMedicine Online; www.rbmonline.com/article/2369 on web 19 October 2006 Case report Successful pregnancy after ICSI with strontium oocyte activation

More information

Unit 6: Study Guide Cell Division. diploid gene allele interphase (G1, S, G2) prophase metaphase anaphase

Unit 6: Study Guide Cell Division. diploid gene allele interphase (G1, S, G2) prophase metaphase anaphase Unit 6: Study Guide Cell Division 1. Define: chromatin chromosome chromatid pair (sister chromatid) centromere spindle fibers haploid diploid gene allele interphase (G1, S, G2) prophase metaphase anaphase

More information

Animal Development. Lecture 3. Germ Cells and Sex

Animal 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 information

Optimal ICSI timing after the first polar body extrusion in in vitro matured human oocytes

Optimal ICSI timing after the first polar body extrusion in in vitro matured human oocytes Human Reproduction Vol.22, No.7 pp. 1991 1995, 2007 Advance Access publication on May 18, 2007 doi:10.1093/humrep/dem124 Optimal ICSI timing after the first polar body extrusion in in vitro matured human

More information

Why do cells reproduce?

Why 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 information

Symposium: Genetic and epigenetic aspects of assisted reproduction Development of artificial gametes

Symposium: Genetic and epigenetic aspects of assisted reproduction Development of artificial gametes RBMOnline - Vol 16. No 4 2008 539-544 Reproductive BioMedicine Online; www.rbmonline.com/article/3205 on web 22 February 2008 Symposium: Genetic and epigenetic aspects of assisted reproduction Development

More information

Abstract. Introduction. RBMOnline - Vol 9. No Reproductive BioMedicine Online; on web 23 June 2004

Abstract. Introduction. RBMOnline - Vol 9. No Reproductive BioMedicine Online;  on web 23 June 2004 RBMOnline - Vol 9. No 2. 2004 210-221 Reproductive BioMedicine Online; www.rbmonline.com/article/1401 on web 23 June 2004 Article Preimplantation genetic diagnosis with HLA matching Dr Svetlana Rechitsky

More information

Article Effect of gonadotrophin priming on in-vitro maturation of oocytes collected from women at risk of OHSS

Article Effect of gonadotrophin priming on in-vitro maturation of oocytes collected from women at risk of OHSS RBMOnline - Vol 13. No 3. 2006 340 348 Reproductive BioMedicine Online; www.rbmonline.com/article/2328 on web 12 June 2006 Article Effect of gonadotrophin priming on in-vitro maturation of oocytes collected

More information

Chapter 14 Cellular Reproduction

Chapter 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 information

Chromosomes & Cell Division

Chromosomes & Cell Division Chromosomes & Cell Division Cell Division The growth and splitting of cells into two new, identical cells called daughter cells. Mitosis Meiosis DNA replicates Parent cell Chromosomes separate Cell division

More information

Articles Impact of parental gonosomal mosaicism detected in peripheral blood on preimplantation embryos

Articles Impact of parental gonosomal mosaicism detected in peripheral blood on preimplantation embryos RBMOnline - Vol 5. No 3. 306 312 Reproductive BioMedicine Online; www.rbmonline.com/article/699 on web 12 September Articles Impact of parental gonosomal mosaicism detected in peripheral blood on preimplantation

More information

Name: Cell division and cancer review

Name: Cell division and cancer review Name: Cell division and cancer review 1. What type of cell undergoes meiosis? Gamete cells or Somatic cells 2. Define homologous chromosomes. 2 chromosomes with similar structure 3. For each of the following

More information

Effect of Bovine Follicular Fluid Added to the Maturation Medium on Sperm Penetration in Pig Oocytes Matured In Vitro

Effect of Bovine Follicular Fluid Added to the Maturation Medium on Sperm Penetration in Pig Oocytes Matured In Vitro Article Effect of Bovine Follicular Fluid Added to the Maturation Medium on Sperm Penetration in Pig Oocytes Matured In Vitro Abstract Naoki ISOBE Research Associate Graduate School for International Development

More information

Mitosis THE CELL CYCLE. In unicellular organisms, division of one cell reproduces the entire organism Multicellular organisms use cell division for..

Mitosis THE CELL CYCLE. In unicellular organisms, division of one cell reproduces the entire organism Multicellular organisms use cell division for.. Mitosis THE CELL CYCLE In unicellular organisms, division of one cell reproduces the entire organism Multicellular organisms use cell division for.. Development from a fertilized cell Growth Repair Cell

More information

The spindle observation and its relationship with fertilization after intracytoplasmic sperm injection in living human oocytes

The spindle observation and its relationship with fertilization after intracytoplasmic sperm injection in living human oocytes FERTILITY AND STERILITY VOL. 75, NO. 2, FEBRUARY 2001 Copyright 2001 American Society for Reproductive Medicine Published by Elsevier Science Inc. Printed on acid-free paper in U.S.A. The spindle observation

More information

5/25/2015. Replication fork. Replication fork. Replication fork. Replication fork

5/25/2015. Replication fork. Replication fork. Replication fork. Replication fork Mutations Chapter 5 Cellular Functions Lecture 3: and Cell Division Most DNA mutations alter the protein product May Make it function better (rarely) Change its function Reduce its function Make it non-functional

More information

To General Embryology Dr: Azza Zaki

To 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 information

Article Impact of meiotic and mitotic non-disjunction on generation of human embryonic stem cell lines

Article Impact of meiotic and mitotic non-disjunction on generation of human embryonic stem cell lines RBMOn - Vol 18. No 1. 2009 120-126 Reproductive BioMedicine On; www.rbmon.com/article/3656 on web 21 November 2008 Article Impact of meiotic and mitotic non-disjunction on generation of human embryonic

More information

Chapter 13 Pre-Test Question 2

Chapter 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 information

Cell Division. Chromosome structure. Made of chromatin (mix of DNA and protein) Only visible during cell division

Cell 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 information

Cell cycle and apoptosis

Cell cycle and apoptosis Cell cycle and apoptosis Cell cycle Definition Stages and steps Cell cycle Interphase (G1/G0, S, and G2) Mitosis (prophase, metaphase, anaphase, telophase, karyokinesis, cytokinesis) Control checkpoints

More information

Influence of genetic factors on the fertilization of mouse ova in vitro

Influence of genetic factors on the fertilization of mouse ova in vitro Influence of genetic factors on the fertilization of mouse ova in vitro El\l=z:\b\l=i:\etaKaleta Department of Genetics and Evolution, Institute of Zoology, Jagellonian University, Krupnicza 50, 30-060

More information

Pre-Test. 4. Inside of cells, nearly all the genes are located on special structures known as microtubules.

Pre-Test. 4. Inside of cells, nearly all the genes are located on special structures known as microtubules. 1 Pre-Test Directions: Answer each question TRUE OR FALSE. 1. Cells in the bodies of multicelled creatures, such as humans, are reproduced in a way that is very similar to the method used by most protozoa.

More information

Submitted April 2, 2003; accepted June 5, KEY WORDS: Fertility; fertilization; oocyte; reproductive techniques. INTRODUCTION

Submitted April 2, 2003; accepted June 5, KEY WORDS: Fertility; fertilization; oocyte; reproductive techniques. INTRODUCTION ( C 2003) Assisted Reproduction Estimation of Second Polar Body Retention Rate After Conventional Insemination and Intracytoplasmic Sperm Injection: In Vitro Observations From more than 5000 Human Oocytes

More information

Biology 4361 Developmental Biology Exam 1 ID#: October 11, 2005

Biology 4361 Developmental Biology Exam 1 ID#: October 11, 2005 Biology 4361 Developmental Biology Name: Key Exam 1 ID#: October 11, 2005 Multiple choice (one point each) 1. Primordial germ cells a. are immortal b. produce polar bodies c. are haploid d. are somatic

More information

Sexual Reproduction and Meiosis

Sexual 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 information

Cell Division and Mitosis

Cell Division and Mitosis Chromatin-Uncoiled DNA during interphase Cell Division and Mitosis Chromosomes-Tightly coiled DNA Chromatid-One half of a duplicated chromosome. Each is identical and called sister chromatids Centromere-The

More information

Chapter 4 The Chromosome Theory of Inheritance

Chapter 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 information

Rejuvenation of Gamete Cells; Past, Present and Future

Rejuvenation of Gamete Cells; Past, Present and Future Rejuvenation of Gamete Cells; Past, Present and Future Denny Sakkas PhD Scientific Director, Boston IVF Waltham, MA, USA Conflict of Interest I have no conflict of interest related to this presentation.

More information

BIOLOGY. Cell Cycle - Mitosis. Outline. Overview: The Key Roles of Cell Division. identical daughter cells. I. Overview II.

BIOLOGY. Cell Cycle - Mitosis. Outline. Overview: The Key Roles of Cell Division. identical daughter cells. I. Overview II. 2 Cell Cycle - Mitosis CAMPBELL BIOLOGY TENTH EDITION Reece Urry Cain Wasserman Minorsky Jackson Outline I. Overview II. Mitotic Phase I. Prophase II. III. Telophase IV. Cytokinesis III. Binary fission

More information

Cellular Reproduction, Part 2: Meiosis Lecture 10 Fall 2008

Cellular Reproduction, Part 2: Meiosis Lecture 10 Fall 2008 Mitosis & 1 Cellular Reproduction, Part 2: Lecture 10 Fall 2008 Mitosis Form of cell division that leads to identical daughter cells with the full complement of DNA Occurs in somatic cells Cells of body

More information

Ploidy and Human Cell Types. Cell Cycle and Mitosis. DNA and Chromosomes. Where It All Began 11/19/2014. Chapter 12 Pg

Ploidy and Human Cell Types. Cell Cycle and Mitosis. DNA and Chromosomes. Where It All Began 11/19/2014. Chapter 12 Pg Ploidy and Human Cell Types Cell Cycle and Mitosis Chapter 12 Pg. 228 245 Cell Types Somatic cells (body cells) have 46 chromosomes, which is the diploid chromosome number. A diploid cell is a cell with

More information

BIOLOGY 4/6/2015. Cell Cycle - Mitosis. Outline. Overview: The Key Roles of Cell Division. identical daughter cells. I. Overview II.

BIOLOGY 4/6/2015. Cell Cycle - Mitosis. Outline. Overview: The Key Roles of Cell Division. identical daughter cells. I. Overview II. 2 Cell Cycle - Mitosis CAMPBELL BIOLOGY TENTH EDITION Reece Urry Cain Wasserman Minorsky Jackson Outline I. Overview II. Mitotic Phase I. Prophase II. III. Telophase IV. Cytokinesis III. Binary fission

More information

3. What law of heredity explains that traits, like texture and color, are inherited independently of each other?

3. What law of heredity explains that traits, like texture and color, are inherited independently of each other? Section 2: Genetics Chapter 11 pg. 308-329 Part 1: Refer to the table of pea plant traits on the right. Then complete the table on the left by filling in the missing information for each cross. 6. What

More information

4. Which of the following cell parts can be found in human cells, but not plant cells? A. mitochondria B. chloroplast C. centrioles D.

4. Which of the following cell parts can be found in human cells, but not plant cells? A. mitochondria B. chloroplast C. centrioles D. http://chss.sd57.bc.ca/~spearce/science_9/practice%20tests/reproductionpracticetest.htm Sc.9 Biology Practise Test 1. Single-celled organisms use cell division for growth. 2. Chromosomes are made of two

More information

Mitosis/Meiosis Simulation Activities

Mitosis/Meiosis Simulation Activities Mitosis/Meiosis Simulation Activities In this simulation, you will demonstrate an understanding of mitosis, meiosis, segregation, independent assortment, and crossing over, all processes involved with

More information

ASSISTED REPRODUCTIVE TECHNOLOGIES (ART)

ASSISTED REPRODUCTIVE TECHNOLOGIES (ART) ASSISTED REPRODUCTIVE TECHNOLOGIES (ART) Dr. Herve Lucas, MD, PhD, Biologist, Andrologist Dr. Taher Elbarbary, MD Gynecologist-Obstetrician Geneva Foundation for Medical Education and research Definitions

More information

meiosis asexual reproduction CHAPTER 9 & 10 The Cell Cycle, Meiosis & Sexual Life Cycles Sexual reproduction mitosis

meiosis asexual reproduction CHAPTER 9 & 10 The Cell Cycle, Meiosis & Sexual Life Cycles Sexual reproduction mitosis meiosis asexual reproduction CHAPTER 9 & 10 The Cell Cycle, Meiosis & Sexual Sexual reproduction Life Cycles mitosis Chromosomes Consists of a long DNA molecule (represents thousands of genes) Also consists

More information

Telomeres in human oocytes. contribution to chromosome (in)stability?

Telomeres in human oocytes. contribution to chromosome (in)stability? Telomeres in human oocytes and embryos: maternal contribution to chromosome (in)stability? Hartshorne GM, Turner S, Rai J, Wong H-P. Warwick Medical School, University of Warwick, UK geraldine.hartshorne@warwick.ac.uk

More information

Diagnostic Techniques to Improve the Assessment of Human IVF Embryos: Genomics and Proteomics

Diagnostic Techniques to Improve the Assessment of Human IVF Embryos: Genomics and Proteomics Diagnostic Techniques to Improve the Assessment of Human IVF Embryos: Genomics and Proteomics Mandy G Katz-Jaffe Introduction A fundamental component of assisted reproductive technologies (ART) is the

More information

General Biology. Overview: The Key Roles of Cell Division The continuity of life is based upon the reproduction of cells, or cell division

General Biology. Overview: The Key Roles of Cell Division The continuity of life is based upon the reproduction of cells, or cell division General Biology Course No: BNG2003" Credits: 3.00 " " " 8. The Cell Cycle Prof. Dr. Klaus Heese Overview: The Key Roles of Cell Division The continuity of life is based upon the reproduction of cells,

More information

General Biology. Overview: The Key Roles of Cell Division. Unicellular organisms

General Biology. Overview: The Key Roles of Cell Division. Unicellular organisms General Biology Course No: BNG2003 Credits: 3.00 8. The Cell Cycle Prof. Dr. Klaus Heese Overview: The Key Roles of Cell Division The continuity of life is based upon the reproduction of cells, or cell

More information

Biology Developmental Biology Spring Quarter Midterm 1 Version A

Biology Developmental Biology Spring Quarter Midterm 1 Version A Biology 411 - Developmental Biology Spring Quarter 2013 Midterm 1 Version A 75 Total Points Open Book Choose 15 out the 20 questions to answer (5 pts each). Only the first 15 questions that are answered

More information

DNA 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 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 information

The Cell Cycle and Mitosis

The Cell Cycle and Mitosis The Key Roles of Cell Division Chapter 1 The Cell Cycle and Mitosis Cell division Cellular reproduc-on An ability of organisms that best dis-nguishes living things from nonliving ma:er Lectures by Chris

More information

Incidence of Chromosomal Abnormalities from a Morphologically Normal Cohort of Embryos in Poor- Prognosis Patients

Incidence of Chromosomal Abnormalities from a Morphologically Normal Cohort of Embryos in Poor- Prognosis Patients Incidence of Chromosomal Abnormalities from a Morphologically Normal Cohort of Embryos in Poor- Prognosis Patients M. C. MAGLI,1 L. GIANAROLI,1,3 S. MUNNE,2 and A. P. FERRARETTI1 Submitted: December 29,

More information

Effect of chromosomal translocations on the development of preimplantation human embryos in vitro

Effect of chromosomal translocations on the development of preimplantation human embryos in vitro FERTILITY AND STERILITY VOL. 74, NO. 4, OCTOBER 2000 Copyright 2000 American Society for Reproductive Medicine Published by Elsevier Science Inc. Printed on acid-free paper in U.S.A.,2 Effect of chromosomal

More information