Concept 12.

2 The mitotic phase alternates with interphase in the cell cycle Phases of the Cell Cycle Mitotic (M) Phase: includes both mitosis and cytokinesis, is usually the shortest part of the cell cycle Interphase: M phase alternates with much longer state called interphase, which often accounts for about 90% of the cycle. A cell that is bout to divide grows and copies chromosomes in prep. Can be divided into subphases o G1 Phase- first gap Cell grows (and continues to grow as it copies its chromosomes (s)) o S Phase- synthesis Chromosomes are duplicated only during S phases Grows more as completes prep. For cell division o G2 Phase- second gap Grows more as it copies its chromosomes *during all three subphases, a cell that will eventually divide grows by producing proteins and cytoplasmic organelles (ex: mitochondria and endoplasmic reticulum) Particular human cell might undergo 1 division in 24 hours ~M phase occupy less than 1 hour ~S phase may occupy 10-12 hours (or about of cycle) ~G1 and G2 phase occupy rest of time Some cells divide very infrequently or not at all: spend time in G1 or G0 phase G2 of Interphase Nuclear envelope encloses nucleus Nucleus contains one or more nucleoli 2 centrosomes have formed by duplication of a single centrosome (centrosome: regions in animal cells that organize the microtubules of the spindle- each centrosome has 2 centrioles) Chromosomes duplicated during S phase cannot be seen individually b/c havent condensed yet Mitosis a. Prophase Chromatin fibers more coiled- to discrete chromosomes Nucleoli disappear Duplicated chromosome appear as 2 identical sister chromatids joined at centromeres (and in some species, joined all along their arms by cohesins) Mitotic spindle begins forming~ the shorter ones that form are called asters (stars) b. Prometaphase Nuclear envelope breaks Microtubules invades nuclear area Chromosomes more condensed Each (of the 2 chromatids) now has a kinetochore (specialized protein structure at centromere) Some microtubules attach to kinetochores- jerking them back and forth

 Nonkinetochore microtubules interact with opposite pole (spindles) c. Metaphase Centrosomes now at opposite poles Chromosomes convene at metaphase plate- chromosomes centromeres lie at the plate For each chromosome, kinetochores of sister chromatids are attached to kinetochore microtubules coming from opposite poles. Centromere= spindle fiber- part of a chromosome that links sister chromatids Spindle fibers attach to centromere via kinetochore Kinetochore= where spindle fiber attaches on chromosome d. Anaphase Shortest stage Begins when cohesin proteins cleaved Allows sister to part Each chromatid becomes full fledge chromosome Liberated daughter chromosome move toward opposite ends (kinetochore microtubules shorten) ~ chromosomes move centromere first Cell elongates as nonkinetochore microtubules lengthen End: 2 ends of cell have equivalent (complete) collections of chromosomes e. Telophase 2 daughter nuclei form nuclear envelops arise nucleoli reappear chromosomes become less condensed remaining spindle microtubules depolymerized mitosis complete Cytokinesis division of cytoplasm usu. Well under way by late telophase animal cells: cleavage furrow : pinches cell in 2

Mitotic Spindle: A closer look Many events of mitosis depend on mitotic spindle: begins to form in cytoplasm during prophase Structure consists of fibers made of microtubules and associated proteins *other microtubules of cytoskeleton partially disassemble to provide the material used to construct the spindle spindle microtubules elongate (polymerize) by incorporating more subunits of tubulin (shorten by depolymerization) Animal cells Assembly of spindle microtubules starts at centrosome: subcellular region containing material that functions throughout the cell cycle to organize the cells microtubules a pair of centrioles located at center of centrosome: but not essential for cell division

During InterphaseSingle centrosome duplicates, forming 2 centrosomes, which remain together near the nucleus. Prophase and prometaphaseTwo centrosomes move apart during prophase and prometaphase ase microtubules grow out from them. (end of prometaphase): 2 centrosomes, one at each pole of the spindle aster: radial array of short microtubules extends from each chromosome Spindle: includes the centrosomes, the spindle microtubules, and asters Kinetochore: each of 2 sister chromatids of a duplicated chromosome has kinetochore: structure of proteins associated with specific sections of chromosomal DNA at each centromere. 2 kinetochores face in opposite directions during prometaphase, some of the spindle microtubules attach to the kinetochores (kinetochore microtubules) When one of a chromosomes kinetochores is captured by microtubules, the chromosome begins to move toward the pole from which those microtubules extend. -stops moving towards one side when microtubules from other side attach to it too -tug of war until it settles at the middle Metaphase Centromeres of all duplicated chromosomes are on a plane midway between the spindles 2 poles: metaphase plate (imaginary) Meanwhile- microtubules that werent attached to kinetochores have been elongating, and by metaphase they overlap and interact with other nonkinetochore microtubules from the opposite pole of the spindle (like overlapped/crossed fingers) - polar microtubules By metaphase, microtubules of the asters have also grown and are in contact with plasma membrane Spindle is now complete Structure of complete spindle correlates well with function Anaphase commences when enzyme separase cleaves cohesins. Chromosomes move toward opposite ends of cell How do kinetochore microtubules function? Two Mechanisms o Kinetochores walk: walk the chromosomes along the microtubules, which depolymerize at their kinetochore ends after motor proteins have passed Pacman mechanism o Reeled In: reeled in by motor proteins at spindle poles; microtubules depolymerize after they pass by these motor proteins

Nonkinetochore microtubules Responsible for elongating the cell during anaphase (from opposite poles) overlap extensively (metaphase) anaphase: region of overlap is reduced- motor proteins attached to the microtubules walk them away from one another (energy: ATP) microtubules push apart spindle poles also pushed apart, elongating the cell. At same time, microtubules lengthen somewhat (addition of tubulin subunits) Cytokinesis: A closer Look Animal: cleavage First sign of cleavage is appearance of cleavage furrow, shallow groove in cell surface near old metaphase plate Cytoplasmic side: contractile ring of actin microfilaments associated with molecules of protein myosin o Actin microfilaments interact with myosin molecules- cause ring to contract Cleavage furrow deepens until cell pinches into 2 Plant No cleavage furrow During telophase, vesicles derived from Golgi apparatus move along microtubules to middle of cell- coalesce to produce cell plate o Cell wall materials carried in o the vesicles collect in cell plate as it grows o Enlarges until surrounding membrane fuses with plasma membrane Binary Fission in Bacteria Division in half- asexual reproduction Bacteria- most genes are carried on single bacterial chromosome circular DNA molecule and associate proteins Process of cell division is initiated when DNA begins to replicate at specific place: origin of replication, producing two origins. As replicating, one origin moves rapidly toward opposite end Replication is complete, bacterium reaches about 2x size Plasma membrane pinches in Several proteins have been identified that play important roles o One resembles eukaryotic actin functions in bacterial chromosome movement during cell division o Another is related to tubulin- helps pinch the plasma Dinoflagellates Chromosomes attach to nuclear envelope (remains intact during cell division) Microtubules pass through nucleus inside cytoplasmic tunnels, reinforcing spatial orientation of nucleus (which then divides in a process reminiscent of bacterial binary fission) Diatoms and some yeasts Nuclear envelope also remains intact during cell division. Microtubules form a spindle within the nucleus

Microtubules separate the chromosomes, and nucleus splits into 2 daughter nuclei

The Evolution of Mitosis Fact that some proteins involved in bacterial fission are related to eukaryotic proteins that function in mitosis supports hypothesis: mitosis evolved from simpler prokaryotic mechanisms of cell reproduction Possible intermediate stages Suggested by 2 unusual types of nuclear division (Dinoflagellates, diatoms, some yeasts) These 2 modes of nuclear division are thought to be cases where ancestral mechanisms have remained relatively unchanged over evolutionary time BOTH TYPES: nuclear envelope remains intact