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Cell cycle
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For the separation of chromosomes that occurs as part of the cell cycle, see mitosis. For the Academic journal, see Cell Cycle (journal).
See also: Cell division
Life cycle of the cell
Onion (Allium) cells in different phases of the cell cycle. Growth in an 'organism' is carefully controlled by regulating the cell cycle.
The cell cycle, or cell-division cycle, is the series of events that take place in a cell leading to its division and duplication of its DNA (DNA replication) to produce two daughter cells. In bacteria, which lack a cell nucleus, the cell cycle is divided into the B, C, and D periods. The B period extends from the end of cell division to the beginning of DNA replication. DNA replication occurs during the C period. The D period refers to the stage between the end of DNA replication and the splitting of the bacterial cell into two daughter cells.[1] In cells with a nucleus, as in eukaryotes, the cell cycle is also divided into three periods: interphase, the mitotic (M) phase, and cytokinesis. During interphase, the cell grows, accumulating nutrients needed for mitosis, preparing it for cell division and duplicating its DNA. During the mitotic phase, the chromosomes separate. During the final stage, cytokinesis, the chromosomes and cytoplasm separate into two new daughter cells. To ensure the proper division of the cell, there are control mechanisms known as cell cycle checkpoints.
The cell-division cycle is a vital process by which a single-celled fertilized egg develops into a mature organism, as well as the process by which hair, skin, blood cells, and some internal organs are renewed. After cell division, each of the daughter cells begin the interphase of a new cycle. Although the various stages of interphase are not usually morphologically distinguishable, each phase of the cell cycle has a distinct set of specialized biochemical processes that prepare the cell for initiation of cell division
Contents
1 Phases
1.1 G0 phase (quiescence)
1.2 Interphase (intermitosis)
1.2.1 G1 phase (First growth phase or Post mitotic gap phase)
1.2.2 S phase (DNA replication)
1.2.3 G2 phase (growth)
1.3 Mitotic phase (chromosome separation)
1.4 Cytokinesis phase (separation of all cell components)
2 Regulation of eukaryotic cell cycle
2.1 Role of cyclins and CDKs
2.1.1 General mechanism of cyclin-CDK interaction
2.1.2 Specific action of cyclin-CDK complexes
2.2 Inhibitors
2.3 Transcriptional regulatory network
2.4 DNA replication and DNA replication origin activity
3 Checkpoints
4 Fluorescence imaging of the cell cycle
5 Role in tumor formation
6 See also
7 References
8 Further reading
9 External links
Phases[edit]
The cell cycle consists of four distinct phases: G1 phase, S phase (synthesis), G2 phase (collectively known as interphase) and M phase (mitosis or meiosis). M phase is itself composed of two tightly coupled processes: karyokinesis, in which the cell's chromosomes are divided, and cytokinesis, in which the cell's cytoplasm divides forming two daughter cells. Activation of each phase is dependent on the proper progression and completion of the previous one. Cells that have temporarily or reversibly stopped dividing are said to have entered a state of quiescence called G0 phase.
Schematic of the cell cycle. Outer ring: I = Interphase, M = Mitosis; inner ring: M = Mitosis, G1 = Gap 1, G2 = Gap 2, S = Synthesis; not in ring: G0 = Gap 0/Resting[2]
State
Phase
Abbreviation
Description
Resting Gap 0 G0 A phase where the cell has left the cycle and has stopped dividing.
Interphase Gap 1 G1 Cells increase in size in Gap 1. The G1 checkpoint control mechanism ensures that everything is ready for DNA synthesis.
Synthesis S DNA replication occurs during this phase.
Gap 2 G2 During the gap between DNA synthesis and mitosis, the cell will continue to grow. The G2 checkpoint control mechanism ensures that everything is ready to enter the M (mitosis) phase and divide.
Cell division Mitosis M Cell growth stops at this stage and cellular energy is focused on the orderly division into two daughter cells. A checkpoint in the middle of mitosis (Metaphase Checkpoint) ensures that the cell is ready to complete cell division.
After cell division, each of the daughter cells begin the interphase of a new cycle. Although the various stages of interphase are not usually morphologically distinguishable, each phase of the cell cycle has a distinct set of specialized biochemical processes that prepare the cell for initiation of cell division.
G0 phase (quiescence)[edit]
Plant cell cycle
Animal cell cycle
G0 is a resting phase where the cell has left the cycle and has stopped dividing. The cell cycle starts with this phase. The word "post-mitotic" is sometimes used to refer to both quiescent and senescent cells. Non-proliferative (non-dividing) cells in multicellular eukaryotes generally enter the quiescent G0 state from G1 and may remain quiescent for long periods of time, possibly indefinitely (as is often the case for neurons). This is very common for cells that are fully differentiated. Cellular senescence occurs in response to DNA damage and external stress and usually constitutes an arrest in G1. Some cells enter the G0 phase semi-permanently and are considered post-mitotic, e.g., some liver, kidney, and stomach cells. Many cells do not enter G0 and continue to divide throughout an organism's life, e.g., epithelial cells.
Cellular senescence is also a state that occurs in response to DNA damage or degradation that would make a cell's progeny nonviable; it is often a biochemical alternative to the self-destruction of such a damaged cell by apoptosis.
Interphase (intermitosis)[edit]
Before a cell can enter cell division, it needs to take in nutrients. All of the preparations are done during interphase. Interphase is a series of changes that takes place in a newly formed cell and its nucleus, before it becomes capable of division again. It is also called preparatory phase or intermitosis. Previously it was called resting stage because there is no apparent activity related to cell division.Typically interphase lasts for at least 91% of the total time required for the cell cycle.
Interphase proceeds in three stages, G1, S, and G2, followed by the cycle of mitosis and cytokinesis. The cell's nuclear DNA contents are duplicated during S phase but may continue till G2 in case of heterochromatin.
G1 phase (First growth phase or Post mitotic gap phase)[edit]
The first phase within interphase, from the end of the previous M phase until the beginning of DNA synthesis, is called G1 (G indicating gap). It is also called the growth phase. During this phase, the biosynthetic activities of the cell, which are considerably slowed down during M phase, resume at a high rate. The duration of G1 is highly variable, even among different cells of the same species.[3] In this phase, the cell increases its supply of proteins, increases the number of organelles (such as mitochondria, ribosomes), and grows in size. In G1 phase, a cell has three options. (1) To continue cell cycle and enter S phase (2) Stop cell cycle and enter G0 phase for undergoing differentiation. (3) Get arrested in G1 phase hence it may enter G0 phase or re-enter cell cycle. The deciding factor is availability of Nitrogens and storage of energy rich compounds at the deciding point called check point (Restriction point). This check point is called the restriction point or START and is regulated by G1/S cyclins, which cause transition from G1 to S phase. Passage through the G1 check point commits the cell to division.
S phase (DNA replication)[edit]
The ensuing S phase starts when DNA synthesis commences; when it is complete, all of the chromosomes have been replicated, i.e., each chromosome has two (sister) chromatids. Thus, during this phase, the amount of DNA in the cell has effectively doubled, though the ploidy of the cell remains the same. Rates of RNA transcription and protein synthesis are very low during this phase. An exception to this is histone production, most of which occurs during the S phase.[4][5][6]
G2 phase (growth)[edit]
G2 phase occurs after DNA replication and is a period of protein synthesis and rapid cell growth to prepare the cell for mitosis. During this phase microtubules begin to reorganize to form a spindle.
Mitotic phase (chromosome separation)[edit]
Main article: Mitosis
The relatively brief M phase consists of nuclear division (karyokinesis). It is a relatively short period of the cell cycle. M phase is complex and highly regulated. The sequence of events is divided into phases, corresponding to the completion of one set of activities and the start of the next. These phases are sequentially known as:
prophase,
prometaphase,
metaphase,
anaphase,
telophase
Mitosis is the process by which a eukaryotic cell separates the chromosomes in its cell nucleus into two identical sets in two nuclei.[7] During the process of mitosis the pairs of chromosomes condense and attach to fibers that pull the sister chromatids to opposite sides of the cell.[8]
Mitosis occurs exclusively in eukaryotic cells, but occurs in different ways in different species. For example, animals undergo an "open" mitosis, where the nuclear envelope breaks down before the chromosomes separate, while fungi such as Aspergillus nidulans and Saccharomyces cerevisiae (yeast) undergo a "closed" mitosis, where chromosomes divide within an intact cell nucleus.[9] Prokaryotic cells, which lack a nucleus, divide by a process called binary fission.
Cytokinesis phase (separation of all cell components)[edit]
Main article: Cytokinesis
Mitosis is immediately followed by cytokinesis, which divides the nuclei, cytoplasm, organelles and cell membrane into two cells containing roughly equal shares of these cellular components. Mitosis and cytokinesis together define the division of the mother cell into two daughter cells, genetically identical to each other and to their parent cell. This accounts for approximately 10% of the cell cycle.
Because cytokinesis usually occurs in conjunction with mitosis, "mitosis" is often used interchangeably with "M phase". However, there are many cells where mitosis and cytokinesis occur separately, forming single cells with multiple nuclei in a process called endoreplication. This occurs most notably among the fungi and slime moulds, but is found in various groups. Even in animals, cytokinesis and mitosis may occur independently, for instance during certain stages of fruit fly embryonic development.[10] Errors in mitosis can either kill a cell through apoptosis or cause mutations that may lead to cancer.
The process of mitosis is complex and highly regulated. The sequence of events is divided into phases, corresponding to the completion of one set of activities and the start of the next. These stages are prophase, prometaphase, metaphase, anaphase and telophase. During the process of mitosis the pairs of chromosomes condense and attach to fibers that pull the sister chromatids to opposite sides of the cell. The cell then divides in cytokinesis, to produce two identical daughter cells.[8]
Regulation of eukaryotic cell cycle[edit]
Regulation of the cell cycle involves processes crucial to the survival of a cell, including the detection and repair of genetic damage as well as the prevention of uncontrolled cell division. The molecular events that control the cell cycle are ordered and directional; that is, each process occurs in a sequential fashion and it is impossible to "reverse" the cycle.
Role of cyclins and CDKs[edit]

Nobel Laureate
Paul Nurse
Nobel Laureate
Tim Hunt
Two key classes of regulatory molecules, cyclins and cyclin-dependent kinases (CDKs), determine a cell's progress through the cell cycle.[11] Leland H. Hartwell, R. Timothy Hunt, and Paul M. Nurse won the 2001 Nobel Prize in Physiology or Medicine for their discovery of these central molecules.[12] Many of the genes encoding cyclins and CDKs are conserved among all eukaryotes, but in general more complex organisms have more elaborate cell cycle control systems that incorporate more individual components. Many of the relevant genes were first identified by studying yeast, especially Saccharomyces cerevisiae;[13] genetic nomenclature in yeast dubs many of these genes cdc (for "cell division cycle") followed by an identifying number, e.g. cdc25 or cdc20.
Cyclins form the regulatory subunits and CDKs the catalytic subunits of an activated heterodimer; cyclins have no catalytic activity and CDKs are inactive in the absence of a partner cyclin. When activated by a bound cyclin, CDKs perform a common biochemical reaction called phosphorylation that activates or inactivates target proteins to orchestrate coordinated entry into the next phase of the cell cycle. Different cyclin-CDK combinations determine the downstream proteins targeted. CDKs are constitutively expressed in cells whereas cyclins are synthesised at specific stages of the cell cycle, in response to various molecular signals.[14]
General mechanism of cyclin-CDK interaction[edit]
Upon receiving a pro-mitotic extracellular signal, G1 cyclin-CDK complexes become active to prepare the cell for S phase, promoting the expression of transcription factors that in turn promote the expression of S cyclins and of enzymes required for DNA replication. The G1 cyclin-CDK complexes also promote the degradation of molecules that function as S phase inhibitors by targeting them for ubiquitination. Once a protein has been ubiquitinated, it is targeted for proteolytic degradation by the proteasome. However, results from a recent study of E2F transcriptional dynamics at the single-cell level argue that the role of G1 cyclin-CDK activities, in particular cyclin D-CDK4/6, is to tune the timing rather than the commitment of cell cycle entry.[15]
Active S cyclin-CDK complexes phosphorylate proteins that make up the pre-replication complexes assembled during G1 phase on DNA replication origins. The phosphorylation serves two purposes: to activate each already-assembled pre-replication complex, and to prevent new complexes from forming. This ensures that every portion of the cell's genome will be replicated once and only once. The reason for prevention of gaps in replication is fairly clear, because daughter cells that are missing all or part of crucial genes will die. However, for reasons related to gene copy number effects, possession of extra copies of certain genes is also deleterious to the daughter cells.
Mitotic cyclin-CDK complexes, which are synthesized but inactivated during S and G2 phases, promote the initiation of mitosis by stimulating downstream proteins involved in chromosome condensation and mitotic spindle assembly. A critical complex activated during this process is a ubiquitin ligase known as the anaphase-promoting complex (APC), which promotes degradation of structural proteins associated with the chromosomal kinetochore. APC also targets the mitotic cyclins for degradation, ensuring that telophase and cytokinesis can proceed.[16]
Specific action of cyclin-CDK complexes[edit]
Cyclin D is the first cyclin produced in the cell cycle, in response to extracellular signals (e.g. growth factors). Cyclin D binds to existing CDK4, forming the active cyclin D-CDK4 complex. Cyclin D-CDK4 complex in turn phosphorylates the retinoblastoma susceptibility protein (Rb). The hyperphosphorylated Rb dissociates from the E2F/DP1/Rb complex (which was bound to the E2F responsive genes, effectively "blocking" them from transcription), activating E2F. Activation of E2F results in transcription of various genes like cyclin E, cyclin A, DNA polymerase, thymidine kinase, etc. Cyclin E thus produced binds to CDK2, forming the cyclin E-CDK2 complex, which pushes the cell from G1 to S phase (G1/S, which initiates the G2/M transition).[17] Cyclin B-cdk1 complex activation causes breakdown of nuclear envelope and initiation of prophase, and subsequently, its deactivation causes the cell to exit mitosis.[14] A quantitative study of E2F transcriptional dynamics at the single-cell level by using engineered fluorescent reporter cells provided a quantitative framework for understanding the control logic of cell cycle entry, challenging the canonical textbook model. Genes that regulate the amplitude of E2F accumulation, such as Myc, determine the commitment in cell cycle and S phase entry. G1 cyclin-CDK activities are not the driver of cell cycle entry. Instead, they primarily tune the timing of E2F increase, thereby modulating the pace of cell cycle progression.[15]
Inhibitors[edit]
Overview of signal transduction pathways involved in apoptosis, also known as "programmed cell death"
Two families of genes, the cip/kip (CDK interacting protein/Kinase inhibitory protein) family and the INK4a/ARF (Inhibitor of Kinase 4/Alternative Reading Frame) family, prevent the progression of the cell cycle. Because these genes are instrumental in prevention of tumor formation, they are known as tumor suppressors.
The cip/kip family includes the genes p21, p27 and p57. They halt cell cycle in G1 phase, by binding to, and inactivating, cyclin-CDK complexes. p21 is activated by p53 (which, in turn, is triggered by DNA damage e.g. due to radiation). p27 is activated by Transforming Growth Factor of β (TGF β), a growth inhibitor.
The INK4a/ARF family includes p16INK4a, which binds to CDK4 and arrests the cell cycle in G1 phase, and p14ARF which prevents p53 degradation.
Synthetic inhibitors of Cdc25 could also be useful for the arrest of cell cycle and therefore be useful as antineoplastic and anticancer agents.[18]
Transcriptional regulatory network[edit]
Current evidence suggests that a semi-autonomous transcriptional network acts in concert with the CDK-cyclin machinery to regulate the cell cycle. Several gene expression studies in Saccharomyces cerevisiae have identified 800–1200 genes that change expression over the course of the cell cycle.[13][19][20] They are transcribed at high levels at specific points in the cell cycle, and remain at lower levels throughout the rest of the cycle. While the set of identified genes differs between studies due to the computational methods and criteria used to identify them, each study indicates that a large portion of yeast genes are temporally regulated.[21]
Many periodically expressed genes are driven by transcription factors that are also periodically expressed. One screen of single-gene knockouts identified 48 transcription factors (about 20% of all non-essential transcription factors) that show cell cycle progression defects.[22] Genome-wide studies using high throughput technologies have identified the transcription factors that bind to the promoters of yeast genes, and correlating these findings with temporal expression patterns have allowed the identification of transcription factors that drive phase-specific gene expression.[19][23] The expression profiles of these transcription factors are driven by the transcription factors that peak in the prior phase, and computational models have shown that a CDK-autonomous network of these transcription factors is sufficient to produce steady-state oscillations in gene expression).[20][24]
Experimental evidence also suggests that gene expression can oscillate with the period seen in dividing wild-type cells independently of the CDK machinery. Orlando et al. used microarrays to measure the expression of a set of 1,271 genes that they identified as periodic in both wild type cells and cells lacking all S-phase and mitotic cyclins (clb1,2,3,4,5,6). Of the 1,271 genes assayed, 882 continued to be expressed in the cyclin-deficient cells at the same time as in the wild type cells, despite the fact that the cyclin-deficient cells arrest at the border between G1 and S phase. However, 833 of the genes assayed changed behavior between the wild type and mutant cells, indicating that these genes are likely directly or indirectly regulated by the CDK-cyclin machinery. Some genes that continued to be expressed on time in the mutant cells were also expressed at different levels in the mutant and wild type cells. These findings suggest that while the transcriptional network may oscillate independently of the CDK-cyclin oscillator, they are coupled in a manner that requires both to ensure the proper timing of cell cycle events.[20] Other work indicates that phosphorylation, a post-translational modification, of cell cycle transcription factors by Cdk1 may alter the localization or activity of the transcription factors in order to tightly control timing of target genes.[22][25][26]
While oscillatory transcription plays a key role in the progression of the yeast cell cycle, the CDK-cyclin machinery operates independently in the early embryonic cell cycle. Before the midblastula transition, zygotic transcription does not occur and all needed proteins, such as the B-type cyclins, are translated from maternally loaded mRNA.[27]
DNA replication and DNA replication origin activity[edit]
Analyses of synchronized cultures of Saccharomyces cerevisiae under conditions that prevent DNA replication initiation without delaying cell cycle progression showed that origin licensing decreases the expression of genes with origins near their 3' ends, revealing that downstream origins can regulate the expression of upstream genes.[28] This confirms previous predictions from mathematical modeling of a global causal coordination between DNA replication origin activity and mRNA expression,[29][30][31] and shows that mathematical modeling of DNA microarray data can be used to correctly predict previously unknown biological modes of regulation.
Checkpoints[edit]
Main article: Cell cycle checkpoint
Cell cycle checkpoints are used by the cell to monitor and regulate the progress of the cell cycle.[32] Checkpoints prevent cell cycle progression at specific points, allowing verification of necessary phase processes and repair of DNA damage. The cell cannot proceed to the next phase until checkpoint requirements have been met. Checkpoints typically consist of a network of regulatory proteins that monitor and dictate the progression of the cell through the different stages of the cell cycle.
There are several checkpoints to ensure that damaged or incomplete DNA is not passed on to daughter cells. Three main checkpoints exist: the G1/S checkpoint, the G2/M checkpoint and the metaphase (mitotic) checkpoint.
G1/S transition is a rate-limiting step in the cell cycle and is also known as restriction point.[14] This is where the cell checks whether it has enough raw materials to fully replicate its DNA (nucleotide bases, DNA synthase, chromatin, etc.). An unhealthy or malnourished cell will get stuck at this checkpoint.
The G2/M checkpoint is where the cell ensures that it has enough cytoplasm and phospholipids for two daughter cells. But sometimes more importantly, it checks to see if it is the right time to replicate. There are some situations where many cells need to all replicate simultaneously (for example, a growing embryo should have a symmetric cell distribution until it reaches the mid-blastula transition). This is done by controlling the G2/M checkpoint.
The metaphase checkpoint is a fairly minor checkpoint, in that once a cell is in metaphase, it has committed to undergoing mitosis. However that's not to say it isn't important. In this checkpoint, the cell checks to ensure that the spindle has formed and that all of the chromosomes are aligned at the spindle equator before anaphase begins.[33]
While these are the three "main" checkpoints, not all cells have to pass through each of these checkpoints in this order to replicate. Many types of cancer are caused by mutations that allow the cells to speed through the various checkpoints or even skip them altogether. Going from S to M to S phase almost consecutively. Because these cells have lost their checkpoints, any DNA mutations that may have occurred are disregarded and passed on to the daughter cells. This is one reason why cancer cells have a tendency to exponentially accrue mutations. Aside from cancer cells, many fully differentiated cell types no longer replicate so they leave the cell cycle and stay in G0 until their death. Thus removing the need for cellular checkpoints. An alternative model of the cell cycle response to DNA damage has also been proposed, known as the postreplication checkpoint.
Checkpoint regulation plays an important role in an organism's development. In sexual reproduction, when egg fertilization occurs, when the sperm binds to the egg, it releases signalling factors that notify the egg that it has been fertilized. Among other things, this induces the now fertilized oocyte to return from its previously dormant, G0, state back into the cell cycle and on to mitotic replication and division.
p53 plays an important role in triggering the control mechanisms at both G1/S and G2/M checkpoints. In addition to p53, checkpoint regulators are being heavily researched for their roles in cancer growth and proliferation.
Fluorescence imaging of the cell cycle[edit]
Fluorescent proteins visualize the cell cycle progression. IFP2.0-hGem(1/110) fluorescence is shown in green and highlights the S/G2/M phases. smURFP-hCdtI(30/120) fluorescence is shown in red and highlights the G0/G1 phases.
Pioneering work by Atsushi Miyawaki and coworkers developed the fluorescent ubiquitination-based cell cycle indicator (FUCCI), which enables fluorescence imaging of the cell cycle. Originally, a green fluorescent protein, mAG, was fused to hGem(1/110) and an orange fluorescent protein (mKO2) was fused to hCdt1(30/120). Note, these fusions are fragments that contain a nuclear localization signal and ubiquitination sites for degradation, but are not functional proteins. The green fluorescent protein is made during the S, G2, or M phase and degraded during the G0 or G1 phase, while the orange fluorescent protein is made during the G0 or G1 phase and destroyed during the S, G2, or M phase.[34] A far-red and near-infrared FUCCI was developed using a cyanobacteria-derived fluorescent protein (smURFP) and a bacteriophytochrome-derived fluorescent protein (movie found at this link).[35]
Role in tumor formation[edit]
A disregulation of the cell cycle components may lead to tumor formation.[36] As mentioned above, when some genes like the cell cycle inhibitors, RB, p53 etc. mutate, they may cause the cell to multiply uncontrollably, forming a tumor. Although the duration of cell cycle in tumor cells is equal to or longer than that of normal cell cycle, the proportion of cells that are in active cell division (versus quiescent cells in G0 phase) in tumors is much higher than that in normal tissue.[citation needed] Thus there is a net increase in cell number as the number of cells that die by apoptosis or senescence remains the same.
The cells which are actively undergoing cell cycle are targeted in cancer therapy as the DNA is relatively exposed during cell division and hence susceptible to damage by drugs or radiation. This fact is made use of in cancer treatment; by a process known as debulking, a significant mass of the tumor is removed which pushes a significant number of the remaining tumor cells from G0 to G1 phase (due to increased availability of nutrients, oxygen, growth factors etc.). Radiation or chemotherapy following the debulking procedure kills these cells which have newly entered the cell cycle.[14]
The fastest cycling mammalian cells in culture, crypt cells in the intestinal epithelium, have a cycle time as short as 9 to 10 hours. Stem cells in resting mouse skin may have a cycle time of more than 200 hours. Most of this difference is due to the varying length of G1, the most variable phase of the cycle. M and S do not vary much.
In general, cells are most radiosensitive in late M and G2 phases and most resistant in late S phase.
For cells with a longer cell cycle time and a significantly long G1 phase, there is a second peak of resistance late in G1.
The pattern of resistance and sensitivity correlates with the level of sulfhydryl compounds in the cell. Sulfhydryls are natural substances that protect cells from radiation damage and tend to be at their highest levels in S and at their lowest near mitosis.
See also[edit]
Cellular model
Synchronous culture – synchronization of cell cultures
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Jump up ^ Ubersax JA, Woodbury EL, Quang PN, Paraz M, Blethrow JD, Shah K, Shokat KM, Morgan DO (October 2003). "Targets of the cyclin-dependent kinase Cdk1". Nature. 425 (6960): 859–64. Bibcode:2003Natur.425..859U. doi:10.1038/nature02062. PMID 14574415.
Jump up ^ Morgan DO (2007). "2–3". The Cell Cycle: Principles of Control. London: New Science Press. p. 18. ISBN 0-9539181-2-2.
Jump up ^ Omberg L, Meyerson JR, Kobayashi K, Drury LS, Diffley JF, Alter O (October 2009). "Global effects of DNA replication and DNA replication origin activity on eukaryotic gene expression". Molecular Systems Biology. 5: 312. doi:10.1038/msb.2009.70. PMC 2779084. PMID 19888207.
Jump up ^ Alter O, Golub GH, Brown PO, Botstein D (2004). Deutscher MP, Black S, Boehmer PE, D'Urso G, Fletcher TM, Huijing F, Marshall A, Pulverer B, Renault B, Rosenblatt JD, Slingerland JM, Whelan WJ, eds. "Novel Genome-Scale Correlation between DNA Replication and RNA Transcription During the Cell Cycle in Yeast is Predicted by Data-Driven Models" (PDF). Miami Nature Biotechnology Winter Symposium: Cell Cycle, Chromosomes and Cancer. Miami Beach, FL: University of Miami School of Medicine, vol. 15 (January 31 – February 4, 2004).
Jump up ^ Alter O, Golub GH (November 2004). "Integrative analysis of genome-scale data by using pseudoinverse projection predicts novel correlation between DNA replication and RNA transcription". Proceedings of the National Academy of Sciences of the United States of America. 101 (47): 16577–82. Bibcode:2004PNAS..10116577A. doi:10.1073/pnas.0406767101. PMC 534520. PMID 15545604.
Jump up ^ Omberg L, Golub GH, Alter O (November 2007). "A tensor higher-order singular value decomposition for integrative analysis of DNA microarray data from different studies". Proceedings of the National Academy of Sciences of the United States of America. 104 (47): 18371–6. Bibcode:2007PNAS..10418371O. doi:10.1073/pnas.0709146104. PMC 2147680. PMID 18003902.
Jump up ^ Elledge SJ (December 1996). "Cell cycle checkpoints: preventing an identity crisis". Science. 274 (5293): 1664–72. Bibcode:1996Sci...274.1664E. doi:10.1126/science.274.5293.1664. PMID 8939848.
Jump up ^ LeMaire-Adkins R, Radke K, Hunt PA (December 1997). "Lack of checkpoint control at the metaphase/anaphase transition: a mechanism of meiotic nondisjunction in mammalian females". The Journal of Cell Biology. 139 (7): 1611–9. doi:10.1083/jcb.139.7.1611. PMC 2132649. PMID 9412457.
Jump up ^ Sakaue-Sawano A, Kurokawa H, Morimura T, Hanyu A, Hama H, Osawa H, Kashiwagi S, Fukami K, Miyata T, Miyoshi H, Imamura T, Ogawa M, Masai H, Miyawaki A (February 2008). "Visualizing spatiotemporal dynamics of multicellular cell-cycle progression". Cell. 132 (3): 487–98. doi:10.1016/j.cell.2007.12.033. PMID 18267078.
Jump up ^ Rodriguez EA, Tran GN, Gross LA, Crisp JL, Shu X, Lin JY, Tsien RY (September 2016). "A far-red fluorescent protein evolved from a cyanobacterial phycobiliprotein". Nature Methods. 13 (9): 763–9. doi:10.1038/nmeth.3935. PMC 5007177. PMID 27479328.
Jump up ^ Champeris Tsaniras S, Kanellakis N, Symeonidou IE, Nikolopoulou P, Lygerou Z, Taraviras S (June 2014). "Licensing of DNA replication, cancer, pluripotency and differentiation: an interlinked world?". Seminars in Cell & Developmental Biology. 30: 174–80. doi:10.1016/j.semcdb.2014.03.013. PMID 24641889.
Further reading[edit]
Morgan DO (2007). The Cell Cycle: Principles of Control. London: Published by New Science Press in association with Oxford University Press. ISBN 0-87893-508-8.
Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2008). "Chapter 17". Molecular Biology of the Cell (5th ed.). New York: Garland Science. ISBN 978-0-8153-4111-6.
Krieger M, Scott MP, Matsudaira PT, Lodish HF, Darnell JE, Zipursky L, Kaiser C, Berk A (2004). Molecular cell biology. New York: W.H. Freeman and CO. ISBN 0-7167-4366-3.
Watson JD, Baker TA, Bell SP, Gann A, Levine M, Losick R (2004). "Chapter 7". Molecular biology of the gene (5th ed.). San Francisco: Pearson/Benjamin Cummings. ISBN 0-8053-4642-2.
External links[edit]
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David Morgan's Seminar: Controlling the Cell Cycle
The cell cycle & Cell death
Transcriptional program of the cell cycle: high-resolution timing
Cell cycle and metabolic cycle regulated transcription in yeast
Cell Cycle Animation 1Lec.com
Cell Cycle
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Science Creative Quarterly's overview of the cell cycle
KEGG – Human Cell Cycle
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text:
Never Give You Up (Gamble & Huff song)
From Wikipedia, the free encyclopedia
Jump to navigation Jump to search
"Never Give You Up"
Single by Jerry Butler
A-side
"Never Give You Up"
B-side
"Beside You"
Released
1968
Format
45RPM Single
Genre
soul
Length
2:56
Label
Mercury
Songwriter(s)
Gamble, Huff, Butler
Jerry Butler singles chronology
"String Module Error: Match not found"
(String Module Error: Match not found) "Never Give You Up"
(1968) "String Module Error: Match not found"
(String Module Error: Match not found)
"Lost" "Never Give You Up" "Hey, Western Union Man"
Never Give You Up was a single by Jerry Butler that reached no. 20 in the Billboard chart in 1968. It was written by Gamble & Huff. Butler was also credited as co-composer. The song has also been covered by Eddie Floyd, Buddy Greco, Chuck Brown and the Soul Searchers, Leonard Julien III and Rod Stewart.
Contents
[hide]
1 Background
2 Other versions
2.1 1960s
2.2 1970s to 1980s
2.3 1990s to 2000s
3 References
4 External links
Background[edit]
The credited composers on the single are Gamble, Huff and Butler. The single was released on Mercury 72798 in April 1968.[1] The song spent 14 weeks on the Hot 100 chart, eventually peaking at no. 20 on July 6, 1968.[2]
Other versions[edit]
1960s[edit]
The Three Midnighters had their version of the song as the B side of their "Chicano Power" single which was released around 1967 - 1968.[3][4] Eddie Floyd covered the song which appeared on his 1968 Stax album, I've Never Found A Girl.[5] A version by Shirley And The Shirelles was recorded. It was included as the B side of single "Go Away And Find Yourself" which was released on Bell in 1969.[6] Buddy Greco recorded a version of the song which appeared as the B side of his single, "Everybody Gets To Go To The Moon", released on Wand Records in 1970.[7] It was also on his Let the Sunshine In album which was previously released on Scepter Records SPS 579 in 1969.[8] A group called The Road had a version on their self-titled album in which was released on Kama Sutra in 1969.[9]
1970s to 1980s[edit]
In 1977, a single by Bonnie Bramlett and Dobie Gray was released on Capricorn 2089 039.[10] Chuck Brown and the Soul Searchers recorded a version which appeared on their 1979 Bustin' Loose album.[11]
1990s to 2000s[edit]
Leonard Julien III recorded the song and it appeared on his Reflections of Soul album which came out on Modeste Records in 2007.[12] Rod Stewart recorded the song as well and it appears on the RCA 88697603432 release of his 2009 Soulbook album.[13] The Black Keys recorded a version which appeared on their Brothers album which was released in 2010.[14] Their version was also used in the Series 3 Episode 4 of the BBC series Luther, which starred Idris Elba.[15] Tony Strong & the Philly Soul recorded a version that was included on their 2012 album "It Ain’t Over".[16]
References[edit]
Jump up ^ 45Cat - Artist: Jerry Butler Never Give You Up, Catalogue: 72798
Jump up ^ Billboard - Jerry Butler | Chart History, Hot 100, Never Give You Up
Jump up ^ 45Cat - Thee Midniters - Discography
Jump up ^ Standard Catalog of American Records, By Dave Thompson - Page 1251 THREE MIDNIGHTERS, 45s, WHITTIER
Jump up ^ Discogs - Eddie Floyd – I've Never Found A Girl
Jump up ^ 45Cat - Shirley And The Shirelles - Discography, US
Jump up ^ 45Cat - Artist: Buddy Greco, Everybody Gets To Go To The Moon Catalogue: WN 10
Jump up ^ Discogs - Buddy Greco – Let The Sunshine In
Jump up ^ Discogs - The Road – The Road
Jump up ^ 45Cat - Artist: Bonnie Bramlett And Dobie Gray Never Give You Up, Catalogue: 2089 039
Jump up ^ Spotify - Never Gonna Give You Up, By Chuck Brown and the Soul Searchers
Jump up ^ Soul Express - Soul Express Album of the Month in August 2008, LEONARD JULIEN III, Reflections of Soul - Ismo Tenkanen, Soul Express Editor
Jump up ^ AllMusic - Rod Stewart, Soulbook, (CD - RCA #88697603432)
Jump up ^ AllMusic - The Black Keys, Never Gonna Give You Up, Composed by Jerry Butler / Kenny Gamble / Leon Huff
Jump up ^ BBC - Luther, Episode 4, Series 3 Episode 4 of 4
Jump up ^ Soul Express, March 2013 - INTRODUCING... TONY STRONG & THE PHILLY SOUL *
External links[edit]
Jerry Butler - Never Gonna Give You Up (1968)
Buddy Greco - Never gonna give you up - A Danny Whitfield Mix
Bonnie Bramlett & Dobie Gray - Never gonna give you up - Danny Whitifield Mix
Retrieved from "https://en.wikipedia.org/w/index.php?title=Never_Give_You_Up_(Gamble_%26_Huff_song)&oldid=824426568"
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Cosmopolitan of Las Vegas
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The Cosmopolitan of Las Vegas
Location
Paradise, Nevada 89109
Address
3708 South Las Vegas Boulevard
Opening date
December 15, 2010; 6 years ago (December 15, 2010)
No. of rooms
3,027
Total gaming space
110,000 sq ft (10,000 m2)
Signature attractions
The Chelsea
The Chandelier
Marquee Nightclub & Dayclub
Rose. Rabbit. Lie.
The Boulevard Pool
P3 Art Studio
Notable restaurants
China Poblano
D.O.C.G.
é
Estiatorio Milos
The Henry
Holsteins
Jaleo
Rose. Rabbit. Lie
Scarpetta
STK
Casino type
Land-based
Owner
The Blackstone Group
Architect
Friedmutter Group (executive architect); Arquitectonica
Coordinates
36°06′36″N 115°10′31″W / 36.1100°N 115.1753°W / 36.1100; -115.1753Coordinates: 36°06′36″N 115°10′31″W / 36.1100°N 115.1753°W / 36.1100; -115.1753
Website
www.cosmopolitanlasvegas.com
[1]
The Cosmopolitan of Las Vegas[2] (commonly referred to simply as The Cosmopolitan or The Cosmo) is a luxury resort casino and hotel on the Las Vegas Strip in Paradise, Nevada. The resort opened on December 15, 2010, and is located just south of the Bellagio on the west side of Las Vegas Boulevard.
It consists of two highrise towers, the Boulevard Tower and the Chelsea Tower, both of which are 184 meters (603 ft) tall.[3] The $3.9 billion project features 3,027 rooms, a 110,000 sq ft (10,000 m2) casino,[4] 300,000 sq ft (28,000 m2) of retail and restaurant space, a 40,000 sq ft (3,700 m2) spa and fitness facility, a 3,200-seat theater, and 150,000 sq ft (14,000 m2) of meeting and convention space.
In 2013, the hotel was rated "The Best Hotel in the World" by Gogobot.[5] In 2015, the resort was named to the Condé Nast Traveller Gold List as one of the "Top Hotels in the World".[6]
Contents
[hide]
1 Amenities
2 History
3 Restaurants
4 Controversy
5 In popular culture
6 Gallery
7 See also
8 References
9 Sources
10 External links
Amenities[edit]
Cosmopolitan features 3,027 hotel rooms[7][8], many of which feature their own private terrace; a 110,000 sq ft (10,000 m2) casino; 300,000 sq ft (28,000 m2) of retail and restaurant space; a 40,000 sq ft (3,700 m2) spa and fitness facility; a 3,200-seat theater; and 150,000 sq ft (14,000 m2) of meeting and convention space. The Cosmopolitan's 100,000 sq ft (9,300 m2) casino features views of the Las Vegas Strip. The Pools at the Cosmopolitan features three different types: a relaxing pool, day club pool and nightclub pool.
The Cosmopolitan is also home to the Marquee Nightclub & Dayclub, which was the top grossing nightclub in the United States in 2012.[9] In January 2014, the Cosmopolitan added a new nightclub concept, Rose. Rabbit. Lie.
History[edit]
Plans for the property were first announced in April 2004.[10] The developer, 3700 Associates, was a joint venture formed by David Friedman (a former Las Vegas Sands executive), Ian Bruce Eichner (a real estate developer), and Soros Fund Management.[10] The developers purchased the site, an 8.5-acre U-shaped parcel surrounding the Jockey Club timeshare building, for $90 million from a company controlled by New Frontier owner Margaret Elardi.[10] Further details about the project, including the Cosmopolitan name, were released in November 2004.[11]
The Cosmopolitan's design team was led by Friedmutter Group as executive architect, with Arquitectonica as the design architect for the building's themed exterior. The building was engineered by DeSimone Consulting Engineers. The interior design team included Digital Kitchen, Prophet, the Friedmutter Group, The Rockwell Group, Jeffrey Beers, Adam Tihany, and Bentel & Bentel.
The resort was built on what used to be the parking lot for the Jockey Club. Because the Cosmopolitan occupies much of the parking lot, it was agreed that the Club residents could use part of the Cosmopolitan's parking garage.[citation needed]
The Cosmopolitan was the second Las Vegas hotel, after The Palazzo, to feature an underground parking garage underneath the hotel. As a result, the parking garage was built first. In December 2007, work finished on the 70-foot (21 m) hole for the parking structure, while other foundation work remained in progress. The hotel was originally planned to open and be operated by Hyatt as the Grand Hyatt Las Vegas.
Original plans called for the casino to be on the second floor, but this was later changed and the casino was built on ground level, like most other Las Vegas hotel-casinos. Planned condo units were cancelled and replaced with studios and other hotel rooms.[12]
In January 2008, it was reported that the $3.9 billion project faced financial complications, as Eichner's company defaulted on a $760 million construction loan from Deutsche Bank when the developer missed a payment after failing to secure refinancing for the project. Construction moved forward as the developers searched for new financing. In late February 2008, Global Hyatt Corporation and New York-based Marathon Asset Management agreed to recapitalize the condominium-hotel project.[13] however, one month later the developer said Deutsche Bank AG would begin foreclosure proceedings.[14] They bought the hotel for $1 billion during the summer and hired The Related Cos., developers of Time Warner Center in New York, to re-position the asset, manage the development process and assist in leasing the retail and restaurant collection.[15][16] Related recommended many revisions, including bringing the casino entrance onto the strip.[17]
In June 2008, Hearst Corp filed a trademark suit against the owners of the casino. Hearst owns the trademark to Cosmopolitan magazine. In March 2010, the suit was settled, and the resort was renamed Cosmopolitan of Las Vegas.[2]
In August 2008, it was announced that MGM Mirage, Starwood Hotels & Resorts Worldwide, Hyatt and Hilton were in talks to acquire the property.[18] It was speculated that MGM Mirage would integrate the project into Bellagio and CityCenter; Starwood were to establish its W and St. Regis brands; and Hyatt would have continued with its plans to operate a Grand Hyatt. In April 2009, the Sun reported that the hotel would be managed by Hilton and would become the Hilton's first in their new Denizen hotel line.[19] Later that month, however, those plans changed; Starwood sued Hilton, claiming trade-secret theft and essentially killing the Denizen brand.[16]
In June 2009, 400 homeowners filed a lawsuit against the developers, claiming breach of contract and seeking refunds for their deposits. They believed that the projected finish date of June 2010 was unrealistic and expressed fear that the developers might turn the condo rooms into hotel rooms only or "finish the building as a shell and not do any interior work."[16]
In April 2010, it was announced that the Cosmopolitan would open in stages, beginning in December and ending in July 2011. It was the only hotel-casino to open on the Strip in 2010. The project officially opened on December 15, 2010, and became part of Marriott International's Autograph Collection, a collection of independent hotels with access to Marriott's reservation and rewards system. In January 2014, the Cosmopolitan announced that points through their Identity rewards program could be redeemed at 3,800 of Marriott's properties.[20] Cosmopolitan is also partnered with the Ritz-Carlton, which is Ritz's first presence on the Las Vegas Strip and their second property in the Las Vegas area.
In May 2014, the Cosmopolitan was sold by Deutsche Bank to Blackstone Group for $1.73 billion.[21]
The July 2015 fire at the Cosmopolitan, seen from the rooftop terrace of Marriott's Grand Chateau. The picture was taken twenty minutes after the fire was reported.
On July 25, 2015, a fire broke out on the pool deck of the resort, burning trees and cabanas, and sending plumes of smoke into the air. Two people were treated for smoke inhalation as a result of the blaze, including one person who was transported to a local hospital.[22] The cause of the fire was not immediately known.[23]
Restaurants[edit]
See also: List of restaurants in the Las Vegas Valley
In March 2010, the casino announced several celebrity chefs and restaurants that would open there.[24] Included were Bruce and Eric Bromberg's Blue Ribbon, Costas Spiliadis' Estiatorio Milos, Scott Conant's Scarpetta, and David Myers' Comme Ça.
On May 2, the resort announced that José Andrés would be joining the resort with three restaurants, creating his namesake restaurant, "é by José Andrés", and one based on his tapas restaurant, Jaleo. Another restaurant, China Poblano, is a new concept combining Mexican and Chinese cuisine.
Additional restaurants include: D.O.C.G. a restaurant and wine bar by Scott Conant; Holstein's, a specialty burger restaurant. Also, STK, a steakhouse by Todd Mark Miller; along with The Henry, Va Bene Caffè, and Wicked Spoon.[25]
Controversy[edit]
In April 2011, Cosmopolitan security staff allegedly removed a transgender guest named Stephanie from a women's restroom, photographed her, and said that she would be banned for life if she did not leave the premises. Shortly after the incident, the hotel-casino was flooded with complaints on its Facebook page, which prompted the hotel-casino to issue an apology to the transgender community and to Stephanie that they would "welcome her back to the resort anytime." The incident also prompted the hotel-casino to train its staff on awareness initiatives involving the sensitive issue.[26]
In popular culture[edit]
The Cosmopolitan was the setting for The Killers music video for "Shot at the Night".[27]
The Cosmopolitan was the setting for Top Chef Masters Season 4.[28]
Gallery[edit]
Night view from the east side.
Cosmopolitan construction as seen from Caesars Palace
The Cosmopolitan nearing completion in March 2010
The street front view from across the Strip
Viewed from the Las Vegas Eiffel Tower
Night view
See also[edit]
Autograph Collection Hotels
List of tallest buildings in Las Vegas
References[edit]
Jump up ^ "Deutsche Bank sells Las Vegas casino to Blackstone for $1.7 bln". Reuters. Retrieved 1 January 2015.
^ Jump up to: a b Arnold M. Knightly (2010-03-20). "Dispute settled, Strip resort renamed". Las Vegas Review-Journal.
Jump up ^ "Las Vegas - The Skyscraper Center". Retrieved October 17, 2016.
Jump up ^ Christopher Palmeri; Beth Jinks (16 April 2014). "Deutsche Bank Said to Seek Sale of Cosmopolitan Resort". Bloomberg. Retrieved 13 May 2014.
Jump up ^ "15 Best Hotels In The World". Huffington Post. November 29, 2012.
Jump up ^ "The Gold List 2016". Retrieved October 17, 2016.
Jump up ^ Moore, Thomas (9 August 2017). "Cosmopolitan launching $100 million-plus room refresh". Las Vegas Sun. Retrieved 9 August 2017.
Jump up ^ "Newly updated guest rooms to feature modern design, bold art and advanced technology". 9 August 2017. Retrieved 20 August 2017.
Jump up ^ Domanick, Andrea (February 21, 2013). "Party capital: Seven of 10 top-grossing U.S. nightclubs are in Las Vegas". Las Vegas Sun.
^ Jump up to: a b c "Details on new Las Vegas Strip development unveiled". Las Vegas Sun. April 14, 2004 – via NewsBank.
Jump up ^ "Partnership names planned Strip property". Las Vegas Sun. November 22, 2004 – via NewsBank.
Jump up ^ Stutz, Howard (April 10, 2010). "Cosmopolitan of Las Vegas sees up-high views as draw". Las Vegas Review-Journal.
Jump up ^ Knightly, Arnold (February 28, 2008). "DEVELOPMENT: Project saved by pair's aid". Las Vegas Review-Journal.
Jump up ^ Forsyth, Jennifer (March 15, 2008). "Foreclosure on Las Vegas Casino to Begin". Wall Street Journal.
Jump up ^ "Buyout keeps Cosmopolitan in play". Las Vegas Sun. 2008-09-04.
^ Jump up to: a b c Illia, Tony. "Homebuyers sue Cosmopolitan". Las Vegas Business Press.
Jump up ^ Berzon, Alexandra (November 17, 2010). "Deutsche Bank Doubles Down With a Casino". The Wall Street Journal.
Jump up ^ Miller, Matt. "MGM Mirage, Hilton Hotels in Talks to Acquire Cosmopolitan Casino". Online Casino Advisory.
Jump up ^ Benston, Liz (March 11, 2009). "New Hilton brand expected to replace Cosmopolitan". Las Vegas Sun.
Jump up ^ Trejos, Nancy (January 23, 2014). "Cosmopolitan guests can now use Marriott points". USA Today.
Jump up ^ "Deutsche Bank sells casino to Blackstone for $1.7 billion". Reuters. Retrieved 1 January 2015.
Jump up ^ "Fire at the Cosmopolitan hotel in Las Vegas injures 2". BNO News. 25 July 2015. Retrieved 25 July 2015.
Jump up ^ "Fire on Las Vegas Strip sends heavy smoke into sky". CBS News. 25 July 2015. Retrieved 25 July 2015.
Jump up ^ [1] Archived June 8, 2011, at the Wayback Machine.
Jump up ^ "Las Vegas Restaurants & Las Vegas Fine Dining at The Cosmopolitan Las Vegas". Cosmopolitanlasvegas.com. Retrieved 1 January 2015.
Jump up ^ "Cosmopolitan responds to backlash over report on banned transgender guest". Las Vegas Sun. 2011-04-27.
Jump up ^ "The Killers Give Lovers a Chance in 'Shot at the Night'". Rolling Stone. 2013-09-27.
Jump up ^ "'Top Chef Masters' cooked at Cosmopolitan". Las Vegas Review-Journal. 2012-07-22.
Sources[edit]
'Cosmo says hello: Guests see model of condo-hotel', Howard Stutz, Las Vegas Review-Journal, October 26, 2005.
External links[edit]
Official website
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Bellagio (hotel and casino)

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The Rockwell Group

values: List of Las Vegas casinos that never opened

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values: Sam's Town Hotel and Gambling Hall, Las Vegas

values: Main Street Station Hotel and Casino and Brewery

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Red Rock Resort Spa and Casino

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Gold Spike Hotel and Casino

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Imperial Palace Hotel and Casino

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Sahara Hotel and Casino

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The Mirage (Las Vegas)

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The Venetian, Las Vegas

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Boulder Strip

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The Orleans Hotel and Casino

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Palms Place

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Aladdin Hotel & Casino

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Marina Hotel and Casino

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values: کازموپالیتن لاس وگاس – Persian

values: The Cosmopolitan Resort & Casino – Indonesian

values: ザ・コスモポリタン – Japanese

values: The Cosmopolitan Resort & Casino – Portuguese

values: 拉斯維加斯大都會酒店 – Chinese

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