Chromosome Segregation: Ndc80 Can Carry the Load

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Chromosome Segregation: Ndc80 Can Carry the Load View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Elsevier - Publisher Connector Current Biology Vol 19 No 10 R404 performance. An absence of What are the implications of these 4. Merker, B. (2000). Synchronous chorusing and human origins. In The Origins of entrainment in such species would animal findings for research on human Music, N.L. Wallin, B. Merker, and S. Brown, raise the question of what else is music and its evolution? The first is that eds. (Cambridge, Mass.: The MIT required for entrainment. we now have animal models to further Press), pp. 315–327. 5. Patel, A.D., Iversen, J.R., Bregman, M.R., and One possibility is the propensity to explore the neural and genetic basis for Schulz, I. (2009). Experimental evidence for engage in joint social action. A recent entrainment. The second illustrates the synchronization to a musical beat in a nonhuman animal. Curr. Biol. 19, study of entrainment in human children fundamental point that evolutionary 827–830. [11] showed that young children find convergence or ‘analogy’ allows us to 6. Schachner, A., Brady, T.F., Pepperberg, I.M., it difficult to entrain to a purely test evolutionary hypotheses, such as and Hauser, M.D. (2009). Entrainment to music requires vocal mimicry: Evidence from non- auditory stimulus (a disembodied the vocal mimicry hypothesis [14]. human animals. Curr. Biol. 19, 831–836. metronomic beat), or to a visible Homologous traits represent a single 7. Fitch, W.T. (2006). The biology and evolution of music: A comparative perspective. Cognition drumbeating robot. They nonetheless evolutionary event, and count 100, 173–215. entrain with a human adult in a socially- statistically as a single datapoint. In 8. Patel, A.D. (2006). Musical rhythm, linguistic engaged game-playing context. contrast, when different clades evolve rhythm, and human evolution. Music Perception 24, 99–104. Perhaps a similar propensity for social the same trait convergently, these 9. Patel, A.D. (2008). Music, Language, and the engagement underlies the apparent constitute statistically independent Brain (New York: Oxford University Press). 10. Janik, V.M., and Slater, P.B. (1997). Vocal capacity for parrots, but not other events [15], allowing us to test learning in mammals. Adv. Stud. Behav. 26, birds, to entrain to a beat? hypotheses about the evolution of 59–99. Parrots are long-lived, group-living human music or language that might 11. Kirschner, S., and Tomasello, M. (2009). Joint drumming: Social context facilitates birds, and their open-ended learning otherwise remain ‘just-so-stories’. synchronization in preschool children. J. Exp. abilities are sometimes employed At both mechanistic and functional Child Psychol. 102, 299–314. 12. Farabaugh, S.M., Linzenbold, A., and to develop vocal ‘badges’ of group levels, then, the discovery of parrot Dooling, R.J. (1994). Vocal plasticity in membership [12]. Although the entrainment provides a rich foundation budgerigars (Melopsittacus undulatus): adaptive functions of parrot for further advances in understanding evidence for social factors in the learning of contact calls. J. Comp. Psychol. 108, 81–92. vocalizations remain poorly the biology and evolution of human 13. Rendell, L., and Whitehead, H. (2001). Culture in understood, they appear to be more music. whales and dolphins. Behav. Brain Sci. 24, 309–324. group-oriented than the mostly 14. Fitch, W.T. (2006). On the biology and evolution individual territorial and courtship of music. Music Perception 24, 85–88. displays that typify songbirds, and this 15. Harvey, P.H., and Pagel, M.D. (1991). The References Comparative Method in Evolutionary Biology may be one factor explaining the 1. Clayton, N.S., Bussey, T.J., and Dickinson, A. (Oxford: Oxford University Press). (2003). Can animals recall the past and plan complete dominance of parrot species for the future? Nat. Rev. Neurosci. 4, in the YouTube sample. This 685–691. School of Psychology, University hypothesis also suggests, given the 2. Kuhl, P.K., and Miller, J.D. (1978). Speech of St. Andrews, St. Andrews, perception by the chinchilla: Identification Scotland, KY6 9JP, UK. capacity of dolphins to engage in functions for synthetic VOT stimuli. J. Acoustic. imitation and joint action [13], that the Soc. Am. 63, 905–917. E-mail: [email protected] 3. Wallin, N.L., Merker, B., and Brown, S. (2000). potential for entrainment in this species The Origins of Music (Cambridge, Mass: The deserves a closer experimental look. MIT Press). DOI: 10.1016/j.cub.2009.04.004 Chromosome Segregation: Ndc80 can directly bind microtubules in vitro Can Carry the Load (reviewed in [1]). In budding yeast, the Ndc80 complex is thought to partner with the Dam1 complex to form kinetochore–microtubule attachments, Dynamic attachments between kinetochores and spindle microtubules are perhaps through direct interaction [2]. required for chromosome bi-orientation in mitosis. A new study provides Dam1 complexes are able to form biophysical insight into how the Ndc80 complex may contribute to the rings and non-ring oligomers on formation of these attachments. microtubules in vitro [3,4], and both forms of assemblages can generate Ajit P. Joglekar1 and the constant gain and loss of tubulin load-bearing attachments to growing Jennifer G. DeLuca2,* subunits at the microtubule plus-ends. and shortening microtubule plus-ends The search for molecular components [5,6]. Although much evidence How cells generate the kinetochore– that serve as load-bearing couplers suggests that Dam1 complexes may microtubule attachments required to between kinetochores and serve as kinetochore–microtubule drive chromosome movements in microtubules is active. Much effort has couplers, no convincing Dam1 mitosis has long puzzled cell biologists. focused on the Ndc80 complex — homologs have surfaced in higher This problem is an interesting one, composed of Ndc80 (Hec1 in humans), eukaryotes. as attachments must be strong to Nuf2, Spc24, and Spc25 — since it is This raises the question: can generate forces for chromosome essential for efficient kinetochore– the Ndc80 complex alone form movements, yet flexible to allow for microtubule attachment in cells and it load-bearing attachments to dynamic Dispatch R405 microtubules? A recent report by Powers and colleagues [7] suggests Ndc80- Ndc80 complex: that it can, and they argue it does so Ndc80 complex- Spc24 using a biased-diffusion mechanism. complex-coated coated Spc25 Hill originally proposed biased bead bead Nuf2 diffusion as a mechanism to explain Bead Hec1 chromosome movement coupled with microtubule depolymerization [8]. This mechanism relies on multiple weak interactions of binding proteins with Diffusion rate = 2.3 ± 0.4µm2/sec the plus-end of a microtubule to produce depolymerization-dependent force. Hill modeled the microtubule- binding site at the kinetochore as a sleeve with regularly spaced weak Tension interaction sites covering its inner applied surface based on the electron microscopy structure of the Bead tracks Bead tracks kinetochore available at the time. depolymerizing polymerizing Thermal diffusion of this sleeve microtubule microtubule (facilitated by weak individual interactions) maintains the most favorable position of the sleeve in relation to the plus-end — one that maximizes the number of interactions Current Biology between the sleeve and the microtubule lattice, minimizing the Figure 1. The Ndc80 complex can form load-bearing attachments to microtubules using free energy of the system. a biased-diffusion-based mechanism. Depolymerization at the microtubule Powers et al. [7] demonstrate that beads coated with Ndc80 complexes (top right) bind and plus-end biases the diffusion of the diffuse rapidly along microtubules (top left). Ndc80-complex-coated beads can track depoly- sleeve, and the sleeve then follows the merizing microtubule plus-ends (bottom left) and, in the presence of tension, can track poly- receding plus-end to attain minimum merizing microtubule plus-ends (bottom right). In the experiments described by Powers et al. [7], beads were coated evenly with Ndc80 complexes; however, only a subset is shown in the free energy. This arrangement can figure for clarity. generate a significant force (approximately 12–13 pN) in a microtubule-depolymerization- a catastrophe were able to persistently competent, on its own, to serve as dependent fashion. Although the track depolymerizing plus-ends over a biased-diffusion-based microtubule specific geometry of the binding site long (and physiologically relevant) coupler at the kinetochore. assumed by Hill may not be present distances (average of 4.8 mm). These How does the Ndc80 complex in cells, biased diffusion of the findings are consistent with a recent interface the microtubule lattice to kinetochore, facilitated by multiple study by McIntosh et al. [9] in which facilitate dynamic coupling? X-ray weak interactions with the microtubule beads coated with purified crystallographic data indicate that lattice, remains a compelling force Caenorhabditis elegans Ndc80 both Hec1 and Nuf2 contain calponin generation mechanism. Experiments complex bound microtubules in vitro, homology (CH) domains, which have carried out by Powers et al. [7] and a subset were able to track been implicated in direct microtubule demonstrate that the Ndc80 complex depolymerizing microtubule ends. In binding [12]. The CH domains in Hec1 meets the requirements of the biased the Powers et al. [7] experiments, when and Nuf2 contain prominent positive diffusion
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