1b), reminiscent of the fibrous network found by transmission EM 1. However, when these chromosomes were imaged by 3D-SIM, we found that the chromosome scaffold was composed of networks of hCAP-E, Topo IIα and KIF4 along the chromatid axes ( Fig. 1a) and exhibited continuously alternating distributions, consistent with previous reports 4, 5, 6. We found that hCAP-E (a condensin subunit), Topo IIα and KIF4 are characteristically localized at the central axis of each chromatid ( Fig. To investigate the localization of chromosomal scaffold proteins, an initial immunofluorescence study of over 100 polyamine-isolated human chromosomes (PA chromosomes) was performed using wide-field microscopy with deconvolution. Taken together, our present studies pave the way for understanding chromosome higher order structure in relation to the structural and functional role of the chromosome scaffold.ĭouble-stranded chromosome scaffold (DCS) revealed by super-resolution microscopy, 3D-SIM Furthermore, we found that the chromosome scaffold could be altered and restored in parallel with changes of chromosome morphology, suggesting the structural role of scaffold proteins contributes to the intrinsic integrity of chromosomes. We also examined the structural role of scaffold proteins by RNAi and found that proper assembly of the scaffold proteins is important for the double-stranded scaffold organization as well as chromosome morphology. Using these new methods, we revealed that the chromosome scaffold exhibited a scaffold protein distribution comprising two main lateral strands within the chromatid axis. FIB/SEM was used in conjunction because this enables direct observation of protein localization inside a chromosome at scanning EM resolution 15. Here, we used 3D-SIM, which has a resolution (~120 nm lateral) that is twice the optical diffraction limit 14, to visualize the localization of each scaffold protein and the structure of the scaffold in metaphase chromosomes. The structural and functional roles of the main scaffold proteins, condensin, Topo IIα and KIF4, have been widely studied 2, 3, 4, 5, 6, 10, 11, 12, 13, but it is unclear how these proteins organize the scaffold and regulate chromosome condensation. Due to these discrepancies, the structure of the chromosome scaffold still remains elusive. In contrast, micromechanical force measurements suggest that chromosomes have no rigid scaffold 9. Broad axial localization of condensin and Topo IIα have also been observed using EM with immunogold labelling 7, 8. However, due to the diffraction limit of light, the detailed distribution patterns of each scaffold protein are still ambiguous. Immunofluorescence has permitted identification of condensin and Topo IIα within the axial regions of mitotic chromosomes, with their alternating distribution patterns leading to a ‘barber pole’ model 6. Initial electron microscopy (EM) observations of histone-depleted chromosomes suggested that the chromosome scaffold is a network structure 1. Various approaches have been tried to elucidate chromosome scaffold structure. The axially-positioned chromosome scaffold of both chromatids mainly comprises non-histone proteins: so-called scaffold proteins, including condensin, topoisomerase IIα (Topo IIα) and kinesin family member 4 (KIF4) 2, 3, 4, 5. The backbone was positioned along the chromosome axes and thus termed the ‘chromosome scaffold’ 1. In the late 1970s, Laemmli and colleagues observed a backbone structure in chromosomes after depletion of histone proteins. However, how chromosomes are organized in mitosis still remains one of the most important enigmas in cell biology. Our model provides new insights into chromosome higher order structure.Ĭhromosome condensation is crucial to ensure the fidelity of chromosome segregation during cell division. We therefore propose a new structural model of the chromosome scaffold that includes twisted double strands, consistent with the physical properties of chromosomal bending flexibility and rigidity. This reversion to the original morphology underscores the role of the scaffold for intrinsic structural integrity of chromosomes. We also find that scaffold protein can adaptably recover its original localization after chromosome reversion in the presence of cations. Here, we use three dimensional-structured illumination microscopy (3D-SIM) and focused ion beam/scanning electron microscopy (FIB/SEM) to reveal the axial distributions of scaffold proteins in metaphase chromosomes comprising two strands. However, the organization and function of the scaffold are still controversial. The most important structural finding has been the presence of a chromosome scaffold composed of non-histone proteins so-called scaffold proteins. Chromosome higher order structure has been an enigma for over a century.
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If you made it this far, raise a glass of whatever and say cheers to all! Cheers to the passionate, cheers to the haters, cheers to the memers, cheers to the contrarians! Cheers to the goof leaguers, wood leaguers and the most sparkling diamonds! Cheers to the quickmatchers, cheers to the Vs AI’rs, cheers to the ppl that play every night, or only twice a year! Cheers to Spazzo and Notbelial and all the lurkers too! Cheers to CCL, the 323r’s, the 6. We couldn’t have done it without you, so thanks, really. All of your participation and energy helped us trudge through our difficult days (it’s still a job after all), guide our decision making, and just generally help us feel like we’re part of something more than just data fields and debug maps. This work was not spawned out of some special DWarner pocket dimension though, it occurred here, in the real (digital) world, with my hundreds of (very talented and handsome) coworkers and most importantly, you.ĭespite what community consensus might state, designers are usually present in forum discussions, watching popular streams (special shoutout to FollowGrubby, CCL, and the crew of Into The Nexus) and of course sharing dank memes. Since then, I’ve touched probably every map, dozens of reworks, several core hero designs and of course several thousand bugs. I started on this project some 8 or so years ago as an assistant technical designer, originally tasked with simple data implementations of the first sets of maps and heroes. For more than 120 years, companies have chosen CSC as their business partner. It's painful to look back and see those freshly separated fragments of 'you' suspended in time, but that's life eh? 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LVL 1 : Zergling > Devouring One (Walker)ĭwelling: Spawning Pool + Growth building for that. Nydus Worm/Canal pretty much works like Inferno's Castle Gate Marketplace + Resource Silo = Defiler Mound (With its crystals being the Resource Silo perhaps?) Could be swapped with Lurker Den and Lurker as a potential new War Machine? ) Overmind or even the Mature Chrysalis would be neato for Grail building.īlacksmith = Baneling Nest ? (Not entirely sure. I agree that the Cerebrate buildings would be the Town Hall-line ups and that Hatchery, Lair and Hive are the Fort, Citadel and Castle. Tilting left or right can change the angle a little. Super Jump (Up + B) – Transforms into a squid and jumps straight up. Tilting left or right changes the direction. Splat Roller (Side + B) – Pushes the Splat Roller, burying any opponents hit by it. Tilting up or down adjusts the angle of the attack. Splattershot (B) – Shoots a stream of ink. 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