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Microtubule Dynamics

微管是高动态的结构性、功能性胞内高速公路。细胞骨架这一术语使人联想到的画面是一种刚性的、不可改变的永久性结构。实际上,细胞骨架是一个极具活力的互联网络,它由三种不同的主要成分组成:微管(聚合微管蛋白二聚体)、微丝(聚合肌动蛋白)和中间纤维。其中,微管是最大的细胞骨架成分。它们几乎在每一个细胞过程中都发挥着作用。 它们赋予原本无定形的细胞以结构稳定性和形态。它们也是一个相互联通的、胞内“高速公路”的主要成分,通过这一“高速公路”,各种细胞携带物可经由分子马达蛋白构成的繁忙网络来往穿梭。在有丝分裂过程中, 针对两个有丝分裂纺锤体之一的微管收缩为细胞分裂提供了必要的动力。

细胞微环境不断变化的性质就要求具备适应能力,尤其是在发育过程中。微管等细胞骨架成分凭借其聚合结构促进了这种适应能力的形成。微管的基本单位是由一个α微管蛋白亚基及一个相应β微管蛋白亚基构成的微管蛋白二聚体。这些二聚体聚合形成约24nm宽的中空管状结构。未经修饰微管蛋白的自然状态是不断变化的,其聚合和解聚速率几乎相等。

诸如Rac和Rho等几种常见的GTP酶家族,通过促进微管蛋白二聚体的GDP/GTP交换,间接推动了微管组装。其他微管稳定剂则采取了更为直接的方法。微管相关蛋白如MAP1、MAP2、MAP4或MAPtau会与聚合微管结合,稳定其聚合形式,从而促进组装和微管生长。大多数MAPs是通过磷酸化激活的,而MAPK (map激酶)磷酸化级联反应又为微管组装的过程加入了一个额外的控制层。相反,细胞却可以通过多种方式来调节微管解聚。促进解聚的直接方式包括多种成分,比如通过与α/β微管蛋白二聚体结合从而防止其聚合的Stathmin蛋白。微管切割酶如剑蛋白能够在其结构中间切断微管,而驱动蛋白I的家族成员如KIF2则会随微管移动,使其“磨损”并解聚。

Microtubule-associated proteins

CLASP1 (Cytoplasmic Linker Associated Protein 1):

CLASPs, such as CLASP1, are nonmotor microtubule-associated proteins that interact with CLIPs (e.g., CLIP170\; MIM 179838). CLASP1 is involved in the regulation of microtubule dynamics at the kinetochore and throughout the spindle (Maiato et al., 2003 [PubMed 12837247]).[supplied by OMIM, Mar 2008].   更多...

MAPT (Microtubule-Associated Protein tau):

This gene encodes the microtubule-associated protein tau (MAPT) whose transcript undergoes complex, regulated alternative splicing, giving rise to several mRNA species. MAPT transcripts are differentially expressed in the nervous system, depending on stage of neuronal maturation and neuron type. MAPT gene mutations have been associated with several neurodegenerative disorders such as...   更多...

CLIP1 (CAP-GLY Domain Containing Linker Protein 1):

CLIP2 (CAP-GLY Domain Containing Linker Protein 2):

Tubulins

Tuba1a - TUBA1A

Microtubules of the eukaryotic cytoskeleton perform essential and diverse functions and are composed of a heterodimer of alpha and beta tubulins. The genes encoding these microtubule constituents belong to the tubulin superfamily, which is composed of six distinct families. Genes from the alpha, beta and gamma tubulin families are found in all eukaryotes. The alpha and beta tubulins represent...   更多...

TUBB (Tubulin, beta):

beta 5-tubulin\; polymerizes to form microtubules\; member of a family of structural proteins [RGD, Feb 2006].   更多...

TUBA4A (Tubulin, alpha 4a):

Microtubules of the eukaryotic cytoskeleton perform essential and diverse functions and are composed of a heterodimer of alpha and beta tubulin. The genes encoding these microtubule constituents are part of the tubulin superfamily, which is composed of six distinct families. Genes from the alpha, beta and gamma tubulin families are found in all eukaryotes. The alpha and beta tubulins represent...   更多...

TUBE1 (Tubulin, epsilon 1):

This gene encodes a member of the tubulin superfamily. This protein localizes to the centriolar sub-distal appendages that are associated with the older of the two centrioles after centrosome duplication. This protein plays a central role in organization of the microtubules during centriole duplication. A pseudogene of this gene is found on chromosome 5.[provided by RefSeq, Jan 2009].   更多...

LOC732582 - Tubulin A:

TUBb6 (Tubulin, beta 6):

Stabilization assembly

CLIP1 (CAP-GLY Domain Containing Linker Protein 1):

The protein encoded by this gene links endocytic vesicles to microtubules. This gene is highly expressed in Reed-Sternberg cells of Hodgkin disease. Several transcript variants encoding different isoforms have been found for this gene. [provided by RefSeq, Oct 2011].   更多...

CDC42 (Cell Division Cycle 42 (GTP Binding Protein, 25kDa)):

The protein encoded by this gene is a small GTPase of the Rho-subfamily, which regulates signaling pathways that control diverse cellular functions including cell morphology, migration, endocytosis and cell cycle progression. This protein is highly similar to Saccharomyces cerevisiae Cdc 42, and is able to complement the yeast cdc42-1 mutant. The product of oncogene Dbl was reported to...   更多...

CLASP1 (Cytoplasmic Linker Associated Protein 1):

CLASPs, such as CLASP1, are nonmotor microtubule-associated proteins that interact with CLIPs (e.g., CLIP170\; MIM 179838). CLASP1 is involved in the regulation of microtubule dynamics at the kinetochore and throughout the spindle (Maiato et al., 2003 [PubMed 12837247]).[supplied by OMIM, Mar 2008].   更多...

CLIP2 (CAP-GLY Domain Containing Linker Protein 2):

ROCK1 (rho-Associated, Coiled-Coil Containing Protein Kinase 1):

ROCK2 (rho-Associated, Coiled-Coil Containing Protein Kinase 2):

Destabilization assembly

FIGNL1 (Fidgetin-Like 1):

May regulate osteoblast proliferation and differentiation (By similarity).   更多...

KIF2C (Kinesin Family Member 2C):

The protein encoded by this gene is a member of kinesin-like protein family. Proteins of this family are microtubule-dependent molecular motors that transport organelles within cells and move chromosomes during cell division. This protein is important for anaphase chromosome segregation and may be required to coordinate the onset of sister centromere separation. [provided by RefSeq, Jul 2008].   更多...

vps4b - VPS4B

The protein encoded by this gene is a member of the AAA protein family (ATPases associated with diverse cellular activities), and is the homolog of the yeast Vps4 protein. In humans, two paralogs of the yeast protein have been identified. The former share a high degree of aa sequence similarity with each other, and also with yeast Vps4 and mouse Skd1 proteins. Mouse Skd1 (suppressor of K+...   更多...

EML2 (Echinoderm Microtubule Associated Protein Like 2):

KATNA1 (Katanin P60 (ATPase Containing) Subunit A 1):

KATNB1 (Katanin P80 (WD Repeat Containing) Subunit B 1):

KIF2B (Kinesin Family Member 2B):

VPS4A (Vacuolar Protein Sorting 4 Homolog A (S. Cerevisiae)):

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