Somatic cell cycles

The eukaryotic somatic cell cycle is defined by a sequential order of tasks a dividing cell has to complete it must replicate its DNA, segregate its chromosomes, grow, and divide. The cell cycle can be divided into four discrete phases. DNA replication is restricted to S phase (DNA synthesis phase), which is preceded by a gap phase called G1 and followed by a gap phase called G2. During mitosis (M phase) the sister chromatids are segregated into two new daughter nuclei and mitosis is completed by the division of the cytoplasm termed cytokinesis (Fig. 1). 

Fig. 10.1. The somatic cell cycle in budding yeast, illustrating the successive phases of the cycle as well as checkpoints (Murray, 1989b).

The somatic cell cycle as a double, cdc2-cdk2 oscillator The cell cycle in yeast and in somatic cells appears to be more complex than in embryonic cells as it is subjected to additional controls linking, for example, the onset of mitosis to the successful completion of DNA replication or to the reaching of a critical cellular size (Cross et al, 1989 ... 

A key problem in the yeast and somatic cell cycles is the ordering of the M and S phases (Nurse, 1994), since mitosis should not occur before DNA replication is completed, while the latter process should not start before cell division, preceded by chromosome segregation, has successfully ended. The proper ordering of the M and S phases is the subject of current investigations (Amon et al., 1994 Hayles et al, 1994 Nurse, 1994), which point to the existenee of mutual inhibitory interactions between the reactions controlling the Gi/S and G2/M transitions. The role of these interactions is to prevent the operation of the cyclin-dependent kinase active in one transition when the other transition is in progress. 

ITowever, most normal somatic cells lack telomerase. Consequently, upon every cycle of cell division when the cell replicates its DNA, about 50-nucleotide portions are lost from the end of each telomere. Thus, over time, the telomeres of somatic cells in animals become shorter and shorter, eventually leading to chromosome instability and cell death. This phenomenon has led some scientists to espouse a telomere theory of aging that implicates telomere shortening as the principal factor in cell, tissue, and even organism aging. Interestingly, cancer cells appear immortal because they continue to reproduce indefinitely. A survey of 20 different tumor types by Geron Corporation of Menlo Park, California, revealed that all contained telomerase activity. 

Reik Is anything known about what happens during nuclear cloning If a somatic nucleus is put back into the egg, are these cell cycle controls disrupted ... 

Do later embryonic cell cycles differ from the somatic ones ... 

The cell cycle is a key process that recurs in a periodic manner. Early cell cycles in amphibian embryos are driven by a mitotic oscillator. This oscillator produces the repetitive activation of the cyclin-dependent kinase cdkl, also known as cdc2 [131]. Cyclin synthesis is sufficient to drive repetitive cell division cycles in amphibian embryonic cells [132]. The period of these relatively simple cell cycles is of the order of 30 min. In somatic cells the cell cycle becomes longer, with durations of up to 24 h or more, owing to the presence of checkpoints that ensure that a cell cycle phase is properly completed before the cell progresses to the next phase. The cell cycle goes successively through the phases Gl, S (DNA replication), G2, and M (mitosis) before a new cycle starts in Gl. After mitosis cells can also enter a quiescent phase GO, from which they enter Gl under mitogenic stimulation. 

The interplay between oscillations and bistability has been addressed in detailed molecular models for the cell cycles of amphibian embryos, yeast and somatic cells [138-141]. The predictions of a detailed model for the cell cycle in yeast were successfully compared with observations of more than a hundred mutants [142]. Other theoretical studies focus on the dynamical properties of particular modules of the cell cycle machinery such as that controlhng the Gl/S transition [143]. 

If the cell cycle in amphibian embryonic cells appears to be driven by a limit cycle oscillator, the question arises as to the precise dynamical nature of more complex cell cycles in yeast and somatic cells. Novak et al. [144] constructed a detailed bifurcation diagram for the yeast cell cycle, piecing together the diagrams obtained as a function of increasing cell mass for the transitions between the successive phases of the cell cycle. In these studies, cell mass plays the role of control parameter a critical mass has to be reached for cell division to occur, provided that it coincides with a surge in cdkl activity which triggers the G2/M transition. 

The frequency of the menstrual/estrous cycle varies among species. For example, monoestrous species (e.g., dogs, cats) exhibit 1-2 cycles per year, whereas polyes-trous species (e.g., rodents, primates) exhibit more frequent cycles of shorter duration. Primordial follicles, each consisting of an oocyte surrounded by a single layer of flattened somatic cells known as granulosa cells, are recruited to become primary follicles, a transition marked by increased oocyte size, formation of a glycoprotein matrix (zona pellucida) around the oocyte, and transformation to cuboidal-shaped... 

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