Motor proteins are a class of molecular motors that are able to move along the surface of a suitable substrate. They convert chemical energy into mechanical work by the hydrolysis of ATP. Flagellar rotation, however, is powered by a proton pump.
The best prominent example of a motor protein is the muscle protein myosin which "motors" the contraction of muscle fibers in animals. Motor proteins are the driving force behind most active transport of proteins and vesicles in the cytoplasm. Kinesins and cytoplasmic dyneins play essential roles in intracellular transport such as axonal transport and in the formation of the spindle apparatus and the separation of the chromosomes during mitosis and meiosis. Axonemal dynein, found in cilia and flagella, is crucial to cell motility, for example in spermatozoa, and fluid transport, for example in trachea.
Diseases associated with motor protein defects
The importance of motor proteins in cells becomes evident when they fail to fulfill their function. For example, kinesin deficiencies have been identified as cause for Charcot-Marie-Tooth disease and some kidney diseases. Dynein deficiencies can lead to chronic infections of the respiratory tract as cilia fail to function without dynein. Defects in muscular myosin predictably cause myopathies, whereas defects in unconventional myosin are the cause for Usher syndrome and deafness.
Cytoskeletal motor proteins
Motor proteins utilizing the cytoskeleton for movement fall into two categories based on their substrates: Actin motors such as myosin move along microfilaments through interaction with actin. Microtubule motors such as dynein and kinesin move along microtubules through interaction with tubulin. There are two basic types of microtubule motors: plus-end motors and minus-end motors, depending on the direction in which they "walk" along the microtubule cables within the cell.
Myosins are actin motors and form myosin complexes consisting of two heavy chains with motor heads and two light chains. Derived from the Greek word for muscle, myosin is the protein responsible for generating muscle contraction. By non-processively walking along actin filaments, many molecules of myosin generate enough force to contract muscle tissue. Myosins are also vital in the process of cell division. They are also involved in cytoplasmic streaming, wherein movement along microfilament networks in the cell allows organelles and cytoplasm to stream in a particular direction. Eighteen different classes of myosins are known.
Genomic representation of myosin motors:
Kinesins are a group of related motor proteins that use a microtubule track in antegrade movement. They are vital to the movement of chromosomes during mitosis and are also responsible for shuttling mitochondria, Golgi bodies, and vesicles within eukaryotic cells. Kinesins typically contain two heavy chains with motor heads which move along microtubules via a pseudo-processive asymmetric walking motion, that can be towards the plus-end or the minus-end, depending on the type of kinesin. Fourteen distinct kinesin families are known, with some additional kinesin-like proteins that cannot be classified into these families.
Genomic representation of kinesin motors:
- Fungi (yeast): 6
- Plants (Arabidopsis thaliana): 61
- Insects (Drosophila melanogaster): 25
- Mammals (human): 45
Dyneins are microtubule motors capable of a retrograde sliding movement. Dynein complexes are much larger and more complex than kinesin and myosin motors. Axonemal dynein facilitates the movement of cilia and flagella, and cytoplasmic dynein facilitates transport of intracellular cargos. Compared to 15 types of axonemal dynein, only two cytoplasmic forms are known.
Genomic representation of dynein motors:
- Fungi (yeast): 1
- Plants (Arabidopsis thaliana): 0
- Insects (Drosophila melanogaster): 13
- Mammals (human): 14-15
In contrast to animals, fungi and non-vascular plants, the cells of flowering plants lack dynein motors. However, they contain a larger number of different kinesins. Many of these plant-specific kinesin groups are specialized for functions during plant cell mitosis. Plant cells differ from animal cells in that they have a cell wall. During mitosis, the new cell wall is built by the formation of a cell plate starting in the center of the cell. This process is facilitated by a phragmoplast, a microtubule array unique to plant cell mitosis. The building of cell plate and ultimately the new cell wall requires kinesin-like motor proteins.
Other molecular motors
Besides the motor proteins above, there are many more types of proteins capable of generating forces and torque in the cell. Many of these molecular motors are ubiquitous in both prokaryotic and eukaryotic cells, although some, such as those involved with cytoskeletal elements or chromatin, are unique to eukaryotes. The motor protein prestin, expressed in mammalian cochlear outer hair cells, produces mechanical amplification in the cochlea. It is a direct voltage-to-force converter, which operates at the microsecond rate and possesses piezoelectric properties.
- ATP synthase
- Molecular motors, for a general discussion of natural and synthetic motor molecules
- Protein dynamics
- Shin'ichi Ishiwata, a scientist who specializes in the study of the chemistry and mechanics of muscles and motor proteins
- Hirokawa N, Tekamura R (2003). "Biochemical and molecular characterization of diseases linked to motor proteins". Trends in Biochemical Sciences 28 (10): 558–565. doi:10.1016/j.tibs.2003.08.006. PMID 14559185.
- Thompson RF, Langford GM (2002). "Myosin superfamily evolutionary history". The Anatomical Record 268 (3): 276–289. doi:10.1002/ar.10160. PMID 12382324.
- Vale RD (2003). "The molecular motor toolbox for intracellular transport". Cell 112 (4): 467–480. doi:10.1016/S0092-8674(03)00111-9. PMID 12600311.
- Miki H, Okada Y, Hirokawa N (2005). "Analysis of the kinesin superfamily: insights into structure and function". Trends in Cell Biology 15 (9): 467–476. doi:10.1016/j.tcb.2005.07.006. PMID 16084724.
- Mallik R, Gross SP (2004). "Molecular motors: strategies to get along". Current Biology 14 (22): R971–R982. doi:10.1016/j.cub.2004.10.046. PMID 15556858.
- Vanstraelen M, Inze D, Geelen D (2006). "Mitosis-specific kinesins in Arabidopsis". Trends in Plant Science 11 (4): 167–175. doi:10.1016/j.tplants.2006.02.004. PMID 16530461.
- Smith LG (2002). "Plant cytokinesis: motoring to the finish". Current Biology 12 (6): R202–R209. doi:10.1016/S0960-9822(02)00751-0. PMID 11909547.
- Abdel-Gany I, Day IS, Simmons PK, Reddy ASN (2005). "Origin and evolution of kinesin-like calmodulin-binding protein". Plant Physiology 138 (3): 1711–1722. doi:10.1104/pp.105.060913. PMC 1176440. PMID 15951483.
- Dallos P, Fakler B (2002). "Prestin, a new type of motor protein.". Nat. Rev. Mol. Cell Biol. 3 (2): 104–11. doi:10.1038/nrm730. PMID 11836512.