有丝分裂（mitosis）
【概述】有丝分裂，又称为间接分裂，由W. Fleming (1882)年首次发现于动物及E. Strasburger（1880）年发现于植物。特点是有纺锤体染色体出现，子染色体被平均分配到子细胞，这种分裂方式普遍见于高等动植物(动物和高等植物)。是真核细胞分裂产生体细胞的过程。
【特点】细胞进行有丝分裂具有周期性。即连续分裂的细胞，从一次分裂完成时开始，到下一次分裂完成时为止，为一个细胞周期。一个细胞周期包括两个阶段：分裂间期和分裂期。
【过程】有丝分裂是一个连续的过程，为了描述方便起见，习惯上按先后顺序划分为间期、前期、中期、后期和末期四个时期，在前期和中期之间有时还划分出一个前中期。
间期 有丝分裂间期分为G1、S、G2三个阶段，其中G1期与G2期进行RNA（即核糖核酸）的复制与有关蛋白质的合成，S期进行DNA的复制。在有丝分裂间期，染色质没有高度螺旋化形成染色体，而是以染色质的形式进行DNA（即脱氧核糖核酸）单链复制。有丝分裂间期是有丝分裂全部过程重要准备过程，是一个重要的基础工作。
前期
自分裂期开始到核膜解体为止的时期。间期细胞进入有丝分裂前期时，核的体积增大，由染色质构成的细染色线逐渐缩短变粗，形成染色体。因为染色体在间期中已经复制，所以每条染色体由两条染色单体组成。核仁在前期的后半渐渐消失。在前期末核膜破裂，于是染色体散于细胞质中。动物细胞有丝分裂前期时靠近核膜有两个中心体。每个中心体由一对中心粒和围绕它们的亮域，称为中心质或中心球所组成。由中心体放射出星体丝，即放射状微管。带有星体丝的两个中心体逐渐分开，移向相对的两极（图1）。这种分开过程推测是由于两个中心体之间的星体丝微管相互作用，更快地增长，结果把两个中心体（两对中心粒）推向两极，而于核膜破裂后终于形成两极之间的纺锤体。
前中期 自核膜破裂起到染色体排列在赤道面上为止。核膜的断片残留于细胞质中，与内质网不易区别，在纺锤体的周围有时可以看到它们。
前中期的主要过程是纺锤体的最终形成和染色体向赤道面的运动。纺锤体有两种类型：一为有星纺锤体，即两极各有一个以一对中心粒为核心的星体，见于绝大多数动物细胞和某些低等植物细胞。一为无星纺锤体。两极无星体，见于高等植物细胞（图2）。
曾经认为有星纺锤体含有三种纺锤丝，即三种微管。一种是星体微管，由星体散射出的微管；二是极微管，是由两极分别向相对一级方向伸展的微管，在赤道区来自两极的极微管互相重叠。现在认为极微管可能是由星体微管伸长形成的。三是着丝点微管，与着丝点联结的微管，亦称着丝点丝或牵引丝。着丝点是在染色体的着丝粒的两侧发育出的结构。有报告说着丝点有使微管蛋白聚合成微管的功能。无星纺锤体只有极微管与着丝点微管。
核膜破裂后染色体分散于细胞质中。每条染色体的两条染色单体其着丝点分别通过着丝点与两极相连。由于极微管和着丝微管之间的相互作用，染色体向赤道面运动。最后各种力达到平衡，染色体乃排列到赤道面上。
中期 从染色体排列到赤道面上，到它们的染色单体开始分向两极之前，这段时间称为中期。有时把前中期也包括在中期之内。中期染色体在赤道面形成所谓赤道板。从一端观察可见这些染色体在赤道面呈放射状排列，这时它们不是静止不动的，而是处于不断摆动的状态。中期染色体浓缩变粗，显示出该物种所特有的数目和形态。因此有丝分裂中期适于做染色体的形态、结构和数目的研究，适于核型分析。
后期每条染色体的两条姊妹染色单体分开并移向两极的时期。分开的染色体称为子染色体。子染色体到达两极时后期结束。染色单体的分开常从着丝点处开始，然后两个染色单体的臂逐渐分开。当它们完全分开后就向相对的两极移动。这种移动的速度依细胞种类而异，大体上在0.2～5微米／分之间。平均速度为 1微米／分。同一细胞内的各条染色体都差不多以同样速度同步地移向两极。子染色体向两极的移动是靠纺锤体的活动实现的。
末期从子染色体到达两极开始至形成两个子细胞为止称为末期。此期的主要过程是子核的形成和细胞体的分裂。子核的形成大体上是经历一个与前期相反的过程。到达两极的子染色体首先解螺旋而轮廓消失，全部子染色体构成一个大染色质块，在其周围集合核膜成分，融合而形成子核的核膜，随着子细胞核的重新组成，核内出现核仁。核仁的形成与特定染色体上的核仁组织区的活动有关。
细胞体的分裂称胞质分裂。动物和某些低等植物细胞的胞质分裂是以缢束或起沟的方式完成的。缢束的动力一般推测是由于赤道区的细胞质周边的微丝收缩的结果。微丝的紧缩使细胞在此区域产生缢束，缢束逐渐加深使细胞体最后一分为二。
高等植物细胞的胞质分裂是靠细胞板的形成。在末期，纺锤丝首先在靠近两极处解体消失，但中间区的纺锤丝保留下来，并且微管增加数量，向周围扩展，形成桶状结构，称为成膜体。与形成成膜体的同时，来自内质网和高尔基器的一些小泡和颗粒成分被运输到赤道区，它们经过改组融合而参加细胞板的形成。细胞板逐渐扩展到原来的细胞壁乃把细胞质一分为二（图3）。细胞板由两层薄膜组成，两层薄膜之间积累果胶质，发育成胞间层，两侧的薄膜积累纤维素，各自发育成子细胞的初生壁。
【细胞有丝分裂记忆口诀】有丝分裂并不难
间前中后末相连
前期：膜仁消失现两体
中期：形定数晰赤道齐
后期：点裂数加均两极
末期：两消两现重开始（动物）
两消两现新壁建（植物）
【动植物的不同】动物细胞有丝分裂的过程,与植物细胞的基本相同.不同的特点是:
1.动物细胞有中心体,在细胞分裂的间期,中心体的两个中心粒各自产生了一个新的中心粒,因而细胞中有两组中心粒.在细胞分裂的过程中,两组中心粒分别移向细胞的两极.在这两组中心粒的周围,发出无数条放射线,两组中心粒之间的星射线形成了纺锤丝.
2.动物细胞分裂末期,细胞的中部并不形成细胞板,而是细胞膜从细胞的中部向内凹陷,最后把细胞缢裂成两部分,每部分都含有一个细胞核.这样,一个细胞就分裂成了两个子细胞
【动植物的相同】动物细胞有丝分裂的过程与植物细胞的分裂过程存在两个个十分重要的相同点：
无论是动物细胞分裂过程还是植物细胞分裂过程都会有染色体的出现和纺锤体的形成。（植物：无星线纺锤体；动物：星线纺锤体）。
【意义】有丝分裂的重要意义，是将亲代细胞的染色体经过复制（实质为DNA的复制）以后，精确地平均分配到两个子细胞中去。由于染色体上有遗传物质DNA，因而在生物的亲代和子代之间保持了遗传性状的稳定性。可见，细胞的有丝分裂对于生物的遗传有重要意义。
英文版介绍(Mitosis)Mitosis is the process in which a eukaryotic cell separates the chromosomes in its cell nucleus, into two identical sets in two daughter nuclei. It is generally followed immediately by cytokinesis, which divides the nuclei, cytoplasm, organelles and cell membrane into two daughter cells containing roughly equal shares of these cellular components. Mitosis and cytokinesis together define the mitotic (M) phase of the cell cycle - the division of the mother cell into two daughter cells, genetically identical to each other and to their parent cell.
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. Prokaryotic cells, which lack a nucleus, divide by a process called binary fission.
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.
Because cytokinesis usually occurs in conjunction with mitosis, "mitosis" is often used interchangeably with "mitotic phase". However, there are many cells where mitosis and cytokinesis occur separately, forming single cells with multiple nuclei. This occurs most notably among the fungi and slime moulds, but is found in various different groups. Even in animals, cytokinesis and mitosis may occur independently, for instance during certain stages of fruit fly embryonic development.Errors in mitosis can either kill a cell through apoptosis or cause mutations that may lead to cancer.
Phases of cell cycle and mitosisInterphase
The mitotic phase is a relatively short period of the cell cycle. It alternates with the much longer interphase, where the cell prepares itself for cell division. Interphase is therefore not part of mitosis. Interphase is divided into three phases, G1 (first gap), S (synthesis), and G2 (second gap). During all three phases, the cell grows by producing proteins and cytoplasmic organelles. However, chromosomes are replicated only during the S phase. Thus, a cell grows (G1), continues to grow as it duplicates its chromosomes (S), grows more and prepares for mitosis (G2), and divides (M).
Preprophase
In plant cells only, prophase is preceded by a pre-prophase stage. In highly vacuolated plant cells, the nucleus has to migrate into the center of the cell before mitosis can begin. This is achieved through the formation of a phragmosome, a transverse sheet of cytoplasm that bisects the cell along the future plane of cell division. In addition to phragmosome formation, preprophase is characterized by the formation of a ring of microtubules and actin filaments (called preprophase band) underneath the plasma membrane around the equatorial plane of the future mitotic spindle. This band marks the position where the cell will eventually divide. The cells of higher plants (such as the flowering plants) lack centrioles: with microtubules forming a spindle on the surface of the nucleus and then being organized into a spindle by the chromosomes themselves, after the nuclear membrane breaks down. The preprophase band disappears during nuclear envelope disassembly and spindle formation in prometaphase.
Prophase
Normally, the genetic material in the nucleus is in a loosely bundled coil called chromatin. At the onset of prophase, chromatin condenses together into a highly ordered structure called a chromosome. Since the genetic material has already been duplicated earlier in S phase, the replicated chromosomes have two sister chromatids, bound together at the centromere by the cohesion complex. Chromosomes are visible at high magnification through a light microscope.
Close to the nucleus are structures called centrosomes, which are made of a pair of centriole. The centrosome is the coordinating center for the cell's microtubules. A cell inherits a single centrosome at cell division, which replicates before a new mitosis begins, giving a pair of centrosomes. The two centrosomes nucleate microtubules (which may be thought of as cellular ropes or poles) to form the spindle by polymerizing soluble tubulin. Molecular motor proteins then push the centrosomes along these microtubules to opposite side of the cell. Although centrosomes help organize microtubule assembly, they are not essential for the formation of the spindle, since they are absent from plants, and centrosomes are not always used in meiosis.
Metaphase
As microtubules find and attach to kinetochores in prometaphase, the centromeres of the chromosomes convene along the metaphase plate or equatorial plane, an imaginary line that is equidistant from the two centrosome poles. This even alignment is due to the counterbalance of the pulling powers generated by the opposing kinetochores, analogous to a tug-of-war between people of equal strength. In certain types of cells, chromosomes do not line up at the metaphase plate and instead move back and forth between the poles randomly, only roughly lining up along the midline. Metaphase comes from the Greek μετα meaning "after."
Because proper chromosome separation requires that every kinetochore be attached to a bundle of microtubules (spindle fibres), it is thought that unattached kinetochores generate a signal to prevent premature progression to anaphase without all chromosomes being aligned. The signal creates the mitotic spindle checkpoint.
Anaphase
When every kinetochore is attached to a cluster of microtubules and the chromosomes have lined up along the metaphase plate, the cell proceeds to anaphase (from the Greek ανα meaning “up,” “against,” “back,” or “re-”).
Two events then occur; First, the proteins that bind sister chromatids together are cleaved, allowing them to separate. These sister chromatids, which have now become distinct sister chromosomes, are pulled apart by shortening kinetochore microtubules and move toward the respective centrosomes to which they are attached. Next, the nonkinetochore microtubules elongate, pushing the centrosomes (and the set of chromosomes to which they are attached) apart to opposite ends of the cell. The force that causes the centrosomes to move towards the ends of the cell is still unknown, although there is a theory that suggests that the rapid assembly and breakdown of microtubules may cause this movement.
These two stages are sometimes called early and late anaphase. Early anaphase is usually defined as the separation of the sister chromatids, while late anaphase is the elongation of the microtubules and the microtubules being pulled farther apart. At the end of anaphase, the cell has succeeded in separating identical copies of the genetic material into two distinct populations.
Telophase
Telophase (from the Greek τελος meaning "end") is a reversal of prophase and prometaphase events. It "cleans up" the after effects of mitosis. At telophase, the nonkinetochore microtubules continue to lengthen, elongating the cell even more. Corresponding sister chromosomes attach at opposite ends of the cell. A new nuclear envelope, using fragments of the parent cell's nuclear membrane, forms around each set of separated sister chromosomes. Both sets of chromosomes, now surrounded by new nuclei, unfold back into chromatin. Mitosis is complete, but cell division is not yet complete.
