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2. Effect of the relative shift between the electron density and temperature pedestal position on the pedestal stability in JET-ILW and comparison with JET-C
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Stefanikova, E., Frassinetti, L., Saarelma, S., Loarte, A., Nunes, I., Garzotti, L., Lomas, P., Rimini, F., Drewelow, P., Kruezi, U., Lomanowski, B., De La Luna, E., Meneses, L., Peterka, M., Viola, B., Giroud, C., Litaudon, Maggi C., Abduallev, X., Abhangi, S., Abreu, M., Afzal, P., Aggarwal, M., Ahlgren, K. M., Ahn, T., J. H., Aho, Mantila, Aiba, L., Airila, N., Albanese, M., Aldred, R., Alegre, V., Alessi, D., Aleynikov, E., Alfier, P., Alberto, Alkseev, Allinson, A., Alper, M., Alves, B., Ambrosino, E., Ambrosino, G., Amicucci, R., Amosov, L., Andersson, Sundã©n, Angelone, E., Anghel, M., Angioni, M., Appel, C., Appelbee, L., Arena, C., Ariola, P., Arnichand, M., Arshad, H., Ash, S., Ashikawa, A., Aslanyan, N., Asunta, V., Auriemma, O., Fulvio, Austin, Avotina, Y., Axton, L., Ayres, M. D., Bacharis, C., Baciero, M., Baiã¡o, A., Bailey, D., Baker, S., Balboa, A., Balden, I., Balshaw, M., Bament, N., Banks, R., Baranov, J. W., Barnard, Y. F., Barnes, M. A., Barnes, D., Barnsley, M., Baron, Wiechec, Barrera, Orte, Baruzzo, L., Matteo, Basiuk, Bassan, V., Bastow, M., Batista, R., Batistoni, A., Baughan, P., Bauvir, R., Baylor, B., Bazylev, L., Beal, B., Beaumont, J., Beckers, P. S., Beckett, M., Becoulet, B., Bekris, A., Beldishevski, N., Bell, M., Belli, K., Bellinger, F., Belonohy, M., Ben, Ayed, Benterman, N., Bergsã¥ker, N. A., Bernardo, H., Bernert, J., Berry, M., Bertalot, M., Besliu, L., Beurskens, C., Bieg, M., Bielecki, B., Biewer, J., Bigi, T., Bã¬lkovã¡, M., Binda, P., Bisoffi, F., Bizarro, A., Bjã¶rkas, J. P. S., Blackburn, C., Blackman, J., Blackman, K., Blanchard, T. R., Blatchford, P., Bobkov, P., Boboc, V., Bodnã¡r, A., Bogar, G., Bolshakova, O., Bolzonella, I., Tommaso, Bonanomi, Bonelli, N., Boom, F., Booth, J., Borba, J., Borodin, D., Borodkina, D., Botrugno, I., Bottereau, A., Boulting, C., Bourdelle, P., Bowden, C., Bower, M., Bowman, C., Boyce, C., Boyd, T., Boyer, C., Bradshaw, H. J., Braic, J. M. A., Bravanec, V., Breizman, R., Bremond, B., Brennan, S., Breton, P. D., Brett, S., Brezinsek, A., Bright, S., Brix, M. D. J., Broeckx, M., Brombin, W., Matteo, Broså‚awski, Brown, A., Brown, D. P. D., Bruno, M., Bucalossi, E., Buch, J., Buchanan, J., Buckley, J., Budny, M. A., Bufferand, R., Bulman, H., Bulmer, M., Bunting, N., Buratti, P., Burckhart, P., Buscarino, A., Busse, A., Butler, A., Bykov, N. K., Byrne, I., Cahyna, J., Calabrã², P., Calvo, G., Camenen, I., Camp, Y., Campling, P., Cane, D. C., Cannas, J., Capel, B., Card, A. J., Cardinali, P. J., Carman, A., Carr, P., Carralero, M., Carraro, D., Carvalho, L., Carvalho, B. B., Carvalho, I., Casson, P., Castaldo, F. J., Catarino, C., Caumont, N., Causa, J., Cavazzana, F., Cave, Ayland, Cavinato, K., Cecconello, M., Ceccuzzi, M., Cecil, S., Cenedese, E., Angelo, Cesario, Challis, R., Chandler, C. D., Chandra, M., Chang, D., Chankin, C. S., Chapman, A., Chapman, I. T., Chernyshova, S. 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D., Thompson, P., Thompson, A., Thompson, C. A., Thorne, V. K., Thornton, L., Thrysã¸e, A., Tigwell, A. S., Tipton, P. A., Tiseanu, N., Tojo, I., Tokitani, H., Tolias, M., Tomeå¡, P., Tonner, M., Towndrow, P., Trimble, M., Tripsky, P., Tsalas, M., Tsavalas, M., Tskhakaya, Jun, Turner, D., Turner, I., Turnyanskiy, M. M., Tvalashvili, M., Tyrrell, G., Uccello, S. G. J., Abidin, Ul, Uljanovs, Z., Ulyatt, J., Urano, D., Uytdenhouwen, H., Vadgama, I., Valcarcel, A. P., Valentinuzzi, D., Valisa, M., Vallejos, Olivares, Valovic, P., Van De Mortel, Van, Eester, Van, Renterghem, Van, Rooij, Varje, G. J., Varoutis, J., Vartanian, S., Vasava, S., Vasilopoulou, K., Vega, T., Verdoolaege, J., Verhoeven, G., Verona, R., Verona, Rinati, Veshchev, G., Vianello, E., Vicente, N., Viezzer, J., Villari, E., Villone, S., Vincenzi, F., Pietro, Vinyar, Viola, I., Vitins, B., Vizvary, A., Vlad, Z., Voitsekhovitch, M., Vondrã¡äek, I., Vora, P., Vu, N., Pires De Sa, Wakeling, W. W., Waldon, B., Walkden, C. W. F., Walker, N., Walker, M., Walsh, R., Wang, M., Wang, E., Warder, N., Warren, S., Waterhouse, R. J., Watkins, J., Watts, N. W., Wauters, C., Weckmann, T., Weiland, A., Weisen, J., Weiszflog, H., Wellstood, M., West, C., Wheatley, A. T., Whetham, M. R., Whitehead, S., Whitehead, A. M., Widdowson, B. D., Wiesen, A. M., Wilkinson, S., Williams, J., Wilson, M., Wilson, A. R., Wilson, D. J., Wilson, H. R., Wischmeier, J., Withenshaw, M., Withycombe, G., Witts, A., Wood, D. M., Wood, D., Woodley, R., Wray, C., Wright, S., Wright, J., J. C., Wu, Wukitch, J., Wynn, S., Xu, A., Yadikin, T., Yanling, D., Yao, W., Yavorskij, L., Yoo, V., Young, M. G., Young, C., Young, D., Young, I. D., Zacks, R., Zagorski, J., Zaitsev, R., Zanino, F. S., Zarins, R., Zastrow, A., Zerbini, K. D., Zhang, M., Zhou, W., Zilli, Y., Zoita, E., Zoletnik, V., Zychor, S., I, Universidad de Sevilla. Departamento de Física Atómica, Molecular y Nuclear, Universidad de Sevilla. RNM138: Física Nuclear Aplicada, JET Contributors, Viola, B., Department of Physics, and Materials Physics
- Subjects
Nuclear and High Energy Physics ,Electron density ,Materials science ,Physics::Instrumentation and Detectors ,Thomson scattering ,education ,114 Physical sciences ,01 natural sciences ,010305 fluids & plasmas ,Pedestal ,ASDEX Upgrade ,Physics::Plasma Physics ,Position (vector) ,0103 physical sciences ,Pedestal position ,pedestal stability ,010306 general physics ,Jet (fluid) ,EUROPED ,JET ,pedestal ,pedestal position ,Pedestal stability ,Plasma ,Condensed Matter Physics ,Electron temperature ,Atomic physics ,physics - Abstract
The electron temperature and density pedestals tend to vary in their relative radial positions, as observed in DIII-D (Beurskens et al 2011 Phys. Plasmas 18 056120) and ASDEX Upgrade (Dunne et al 2017 Plasma Phys. Control. Fusion 59 14017). This so-called relative shift has an impact on the pedestal magnetohydrodynamic (MHD) stability and hence on the pedestal height (Osborne et al 2015 Nucl. Fusion 55 063018). The present work studies the effect of the relative shift on pedestal stability of JET ITER-like wall (JET-ILW) baseline low triangularity (δ) unseeded plasmas, and similar JET-C discharges. As shown in this paper, the increase of the pedestal relative shift is correlated with the reduction of the normalized pressure gradient, therefore playing a strong role in pedestal stability. Furthermore, JET-ILW tends to have a larger relative shift compared to JET carbon wall (JET-C), suggesting a possible role of the plasma facing materials in affecting the density profile location. Experimental results are then compared with stability analysis performed in terms of the peeling-ballooning model and with pedestal predictive model EUROPED (Saarelma et al 2017 Plasma Phys. Control. Fusion). Stability analysis is consistent with the experimental findings, showing an improvement of the pedestal stability, when the relative shift is reduced. This has been ascribed mainly to the increase of the edge bootstrap current, and to minor effects related to the increase of the pedestal pressure gradient and narrowing of the pedestal pressure width. Pedestal predictive model EUROPED shows a qualitative agreement with experiment, especially for low values of the relative shift. EURATOM 633053 Swedish Energy Agency 40146-1
- Published
- 2018
3. Corrigendum
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Colas I, Macaulay M, Higgins JD, Phillips D, Barakate A, Posch M, Armstrong SJ, Franklin FC, Halpin C, Waugh R, Ramsay L
- Published
- 2018
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4. Erosion and deposition in the JET divertor during the second ITER-like wall campaign
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Mayer, M., Krat, S., Baron-Wiechec, A., Gasparyan, Y., Heinola, K., Koivuranta, S., Likonen, J., Ruset, C., De Saint-Aubin, G., Litaudon, Widdowson A., Abduallev, X., Abhangi, S., Abreu, M., Afzal, P., Aggarwal, M., Ahlgren, K. M., Ahn, T., J. H., Aho, Mantila, Aiba, L., Airila, N., Albanese, M., Aldred, R., Alegre, V., Alessi, D., Aleynikov, E., Alfier, P., Alberto, Alkseev, Allinson, A., Alper, M., Alves, B., Ambrosino, E., Ambrosino, G., Amicucci, R., Amosov, L., Andersson, Sundã©n, Angelone, E., Anghel, M., Angioni, M., Appel, C., Appelbee, L., Arena, C., Ariola, P., Arnichand, M., Arshad, H., Ash, S., Ashikawa, A., Aslanyan, N., Asunta, V., Auriemma, O., Fulvio, Austin, Avotina, Y., Axton, L., Ayres, M. D., Bacharis, C., Baciero, M., Baiã¡o, A., Bailey, D., Baker, S., Balboa, A., Balden, I., Balshaw, M., Bament, N., Banks, R., Baranov, J. W., Barnard, Y. F., Barnes, M. A., Barnes, D., Barnsley, M., Baron, Wiechec, Barrera, Orte, Baruzzo, L., Matteo, Basiuk, Bassan, V., Bastow, M., Batista, R., Batistoni, A., Baughan, P., Bauvir, R., Baylor, B., Bazylev, L., Beal, B., Beaumont, J., Beckers, P. S., Beckett, M., Becoulet, B., Bekris, A., Beldishevski, N., Bell, M., Belli, K., Bellinger, F., Belonohy, M., Ben, Ayed, Benterman, N., Bergsã¥ker, N. A., Bernardo, H., Bernert, J., Berry, M., Bertalot, M., Besliu, L., Beurskens, C., Bieg, M., Bielecki, B., Biewer, J., Bigi, T., Bã¬lkovã¡, M., Binda, P., Bisoffi, F., Bizarro, A., Bjã¶rkas, J. P. S., Blackburn, C., Blackman, J., Blackman, K., Blanchard, T. R., Blatchford, P., Bobkov, P., Boboc, V., Bodnã¡r, A., Bogar, G., Bolshakova, O., Bolzonella, I., Tommaso, Bonanomi, Bonelli, N., Boom, F., Booth, J., Borba, J., Borodin, D., Borodkina, D., Botrugno, I., Bottereau, A., Boulting, C., Bourdelle, P., Bowden, C., Bower, M., Bowman, C., Boyce, C., Boyd, T., Boyer, C., Bradshaw, H. J., Braic, J. M. A., Bravanec, V., Breizman, R., Bremond, B., Brennan, S., Breton, P. D., Brett, S., Brezinsek, A., Bright, S., Brix, M. D. J., Broeckx, M., Brombin, W., Matteo, Broså‚awski, Brown, A., Brown, D. P. D., Bruno, M., Bucalossi, E., Buch, J., Buchanan, J., Buckley, J., Budny, M. A., Bufferand, R., Bulman, H., Bulmer, M., Bunting, N., Buratti, P., Burckhart, P., Buscarino, A., Busse, A., Butler, A., Bykov, N. K., Byrne, I., Cahyna, J., Calabrã², P., Calvo, G., Camenen, I., Camp, Y., Campling, P., Cane, D. C., Cannas, J., Capel, B., Card, A. J., Cardinali, P. J., Carman, A., Carr, P., Carralero, M., Carraro, D., Carvalho, L., Carvalho, B. B., Carvalho, I., Casson, P., Castaldo, F. J., Catarino, C., Caumont, N., Causa, J., Cavazzana, F., Cave, Ayland, Cavinato, K., Cecconello, M., Ceccuzzi, M., Cecil, S., Cenedese, E., Angelo, Cesario, Challis, R., Chandler, C. D., Chandra, M., Chang, D., Chankin, C. S., Chapman, A., Chapman, I. T., Chernyshova, S. C., Chitarin, M., Giuseppe, Ciraolo, Ciric, G., Citrin, D., Clairet, J., Clark, F., Clark, E., Clarkson, M., Clatworthy, R., Clements, D., Cleverly, C., Coad, M., Coates, J. P., Cobalt, P. A., Coccorese, A., Cocilovo, V., Coda, V., Coelho, S., Coenen, R., Coffey, J. W., Colas, I., Collins, L., Conka, S., Conroy, D., Conway, S., Coombs, N., Cooper, D., Corradino, S. R., Corre, C., Corrigan, Y., Cortes, G., Coster, S., Couchman, D., Cox, A. S., Craciunescu, M. P., Cramp, T., Craven, S., Crisanti, R., Croci, F., Croft, G., Crombã©, D., Crowe, K., Cruz, R., Cseh, N., Cufar, G., Cullen, A., Curuia, A., Czarnecka, M., Dabirikhah, A., Dalgliesh, H., Dalley, P., Dankowski, S., Darrow, J., Davies, D., Davis, O., Day, W., Day, C., I. E., Bock, De, Castro, De, De La Cal, De La Luna, Masi, De, Pablos, De, J. L., Temmerman, De, Tommasi, De, Vries, De, Deakin, P., Deane, K., Degli, Agostini, Dejarnac, F., Delabie, R., Den, Harder, Dendy, N., Denis, R. O., Denner, J., Devaux, P., Devynck, S., Maio, Di, Siena, Di, Troia, Di, Dinca, C., D'Inca, P., Ding, R., Dittmar, B., Doerk, T., Doerner, H., Donnã©, R. P., Dorling, T., S. E., Dormido, Canto, Doswon, S., Douai, S., Doyle, D., Drenik, P. T., Drewelow, A., Drews, P., Duckworth, P., Dumont, P. h., Dumortier, R., Dunai, P., Dunne, D., Äžuran, M., Durodiã©, I., Dutta, F., Duval, P., Dux, B. P., Dylst, R., Dzysiuk, K., Edappala, N., Edmond, P. V., Edwards, J., Edwards, A. M., Eich, J., Ekedahl, T. h., Jorf, El, Elsmore, R., Enachescu, C. G., Ericsson, M., Eriksson, G., Eriksson, F., Eriksson, J., Esposito, L. G., Esquembri, B., Esser, S., Esteve, H. G., Evans, D., Evans, B., Evison, G. E., Ewart, G., Fagan, G. D., Faitsch, D., Falie, M., Fanni, D., Fasoli, A., Faustin, A., Fawlk, J. M., Fazendeiro, N., Fedorczak, L., Felton, N., Fenton, R. C., Fernades, K., Fernandes, A., Ferreira, H., Fessey, J., Fã©vrier, J. A., Ficker, O., Field, O., Fietz, A., Figueiredo, S., Figueiredo, A., Fil, J., Finburg, A., Firdaouss, P., Fischer, M., Fittill, U., Fitzgerald, L., Flammini, M., Flanagan, D., Fleming, J., Flinders, C., Fonnesu, K., Fontdecaba, N., Formisano, J. M., Forsythe, A., Fortuna, L., Fortuna, Zalesna, Fortune, E., Foster, M., Franke, S., Franklin, T., Frasca, T., Frassinetti, M., Freisinger, L., Fresa, M., Frigione, R., Fuchs, D., Fuller, V., Futatani, D., Fyvie, S., Gã¡l, J., Galassi, K., Gaå‚azka, D., Galdon, Quiroga, Gallagher, J., Gallart, J., Galvã¡o, D., Gao, R., Gao, X., Garcia, Y., Garcia, Carrasco, Garcã¬a, Muã±oz, Gardarein, M., Garzotti, J. L., Gaudio, L., Gauthier, P., Gear, E., Gee, D. F., Geiger, S. J., Gelfusa, B., Gerasimov, M., Gervasini, S., Gethins, G., Ghani, M., Ghate, Z., Gherendi, M., Giacalone, M., Giacomelli, J. C., Gibson, L., Giegerich, C. S., Gil, T., Gil, C., Gilligan, L., Gin, S., Giovannozzi, D., Girardo, E., Giroud, J. B., Giruzzi, C., Gerardo, Glã¶ggler, Godwin, S., Goff, J., Gohil, J., Goloborod'Ko, P., Gomes, V., Goncalves, R., Goniche, B., Goodliffe, M., Goodyear, M., Gorini, A., Gosk, G., Goulding, M., Goussarov, R., Gowland, A., Graham, R., Graham, B., Graves, M. E., Grazier, J. P., Grazier, N., Green, P., Greuner, N. R., Grierson, H., Griph, B., Grisolia, F. S., Grist, C., Groth, D., Grove, M., Grundy, R., Grzonka, C. N., Guard, J., Guã©rard, D., Guillemaut, C., Guirlet, C., Gurl, R., Utoh, C., Hackett, H. H., Hacquin, L. J., Hagar, S., Hager, A., Hakola, R., Halitovs, A., Hall, M., S. J., Hallworth, Cook, S. P., Hamlyn, Harris, Hammond, C., Harrington, K., Harrison, C., Harting, J., Hasenbeck, D., Hatano, F., Hatch, Y., Haupt, D. R., Hawes, T. D. V., Hawkes, J., Hawkins, N. C., Hawkins, J., Haydon, P., Hayter, P. W., Hazel, N., Heesterman, S., Heinola, P. J. L., Hellesen, K., Hellsten, C., Helou, T., Hemming, W., Hender, O. N., Henderson, T. C., Henderson, M., Henriques, S. S., Hepple, R., Hermon, D., Hertout, G., Hidalgo, P., Highcock, C., Hill, E. G., Hillairet, M., Hillesheim, J., Hillis, J., Hizanidis, D., Hjalmarsson, K., Hobirk, A., Hodille, J., Hogben, E., Hogeweij, C. H. A., Hollingsworth, G. M. D., Hollis, A., Homfray, S., Horã¡äek, D. A., Hornung, J., Horton, G., Horton, A. R., Horvath, L. D., Hotchin, L., Hough, S. P., Howarth, M. R., Hubbard, P. J., Huber, A., Huddleston, V., Hughes, T. M., Huijsmans, M., Hunter, G. T. A., Huynh, C. L., Hynes, P., Iglesias, A. M., Imazawa, D., Imbeaux, N., Imrã¬å¡ek, F., Incelli, M., Innocente, M., Irishkin, P., Ivanova, Stanik, Jachmich, I., Jacobsen, S., Jacquet, A. S., Jansons, P., Jardin, J., Jã¤rvinen, A., Jaulmes, A., Jednorã³g, F., Jenkins, S., Jeong, I., Jepu, C., Joffrin, I., Johnson, E., Johnson, R., Johnston, T., Jane, Joita, Jones, L., Jones, G., Hoshino, T. T. C., Kallenbach, K. K., Kamiya, A., Kaniewski, K., Kantor, J., Kappatou, A., Karhunen, A., Karkinsky, J., Karnowska, D., Kaufman, I., Kaveney, M., Kazakov, G., Kazantzidis, Y., Keeling, V., Keenan, D. L., Keep, T., Kempenaars, J., Kennedy, M., Kenny, C., Kent, D., Kent, J., Khilkevich, O. N., Kim, E., Kim, H. T., Kinch, H. S., King, A., King, C., King, D., Kinna, R. F., Kiptily, D. J., Kirk, V., Kirov, A., Kirschner, K., Kizane, A., Klepper, G., Klix, C., Knight, A., Knipe, P., Knott, S. J., Kobuchi, S., Kã¶chl, T., Kocsis, F., Kodeli, G., Kogan, I., Kogut, L., Koivuranta, D., Kominis, S., Kã¶ppen, Y., Kos, M., Koskela, B., Koslowski, T., Koubiti, H. R., Kovari, M., Kowalska, Strzè©ciwilk, Krasilnikov, E., Krasilnikov, A., Krawczyk, V., Kresina, N., Krieger, M., Krivska, K., Kruezi, A., Ksiaå¼ek, U., Kukushkin, I., Kundu, A., Kurki, Suonio, Kwak, T., Kwiatkowski, S., Kwon, R., Laguardia, O. J., Lahtinen, L., Laing, A., Lam, A., Lambertz, N., Lane, H. T., Lang, C., Lanthaler, P. T., Lapins, S., Lasa, J., Last, A., Åaszyå„ska, J. R., Lawless, E., Lawson, R., Lawson, A., Lazaros, K. D., Lazzaro, A., Leddy, E., Lee, J., Lefebvre, S., Leggate, X., Lehmann, H. J., Lehnen, J., Leichtle, M., Leichuer, D., Leipold, P., Lengar, F., Lennholm, I., Lerche, M., Lescinskis, E., Lesnoj, A., Letellier, S., Leyland, E., Leysen, M., Li, W., Liang, L., Likonen, Y., Linke, J., Linsmeier, J., Lipschultz, C. h., Liu, B., Liu, G., Schiavo, Lo, Loarer, V. P., Loarte, T., Lobel, A., Lomanowski, R. C., Lomas, B., Lã¶nnroth, P. J., Lã³pez, J., J. M., Lã³pez, Razola, Lorenzini, J., Losada, R., Lovell, U., Loving, J. J., Lowry, A. B., Luce, C., Lucock, T., Lukin, R. M. A., Luna, A., Lungaroni, C., Lungu, M., Lungu, C. P., Lunniss, M., Lupelli, A., Lyssoivan, I., Macdonald, A., Macheta, N., Maczewa, P., Magesh, K., Maget, B., Maggi, P., Maier, C., Mailloux, H., Makkonen, J., Makwana, T., Malaquias, R., Malizia, A., Manas, A., Manning, P., Manso, A., Mantica, M. E., Mantsinen, P., Manzanares, M., Maquet, A., Marandet, P. h., Marcenko, Y., Marchetto, N., Marchuk, C., Marinelli, O., Marinucci, M., Markoviä, M., Marocco, T., Marot, D., Marren, L., Marshal, C. A., Martin, R., Martin, A., Martìn De Aguilera, Martã¬nez, A., F. J., Martã¬n, Solã¬s, Martynova, J. R., Maruyama, Y., Masiello, S., Maslov, A., Matejcik, M., Mattei, S., Matthews, M., Maviglia, G. F., Mayer, F., Mayoral, M., M. L., May, Smith, Mazon, T., Mazzotta, D., Mcadams, C., Mccarthy, R., Mcclements, P. J., Mccormack, K. G., Mccullen, O., Mcdonald, P. A., Mcintosh, D., Mckean, S., Mckehon, R., Meadows, J., Meakins, R. C., Medina, A., Medland, F., Medley, M., Meigh, S., Meigs, S., Meisl, A. G., Meitner, G., Meneses, S., Menmuir, L., Mergia, S., Merrigan, K., Mertens, I. R., Meshchaninov, P. h., Messiaen, S., Meyer, A., Mianowski, H., Michling, S., Middleton, Gear, Miettunen, D., Militello, J., Militello, Asp, Miloshevsky, E., Mink, G., Minucci, F., Miyoshi, S., Mlynã¡å™, Y., Molina, J., Monakhov, D., Moneti, I., Mooney, M., Moradi, R., Mordijck, S., Moreira, S., Moreno, L., Moro, R., Morris, F., Morris, A. W., Moser, J., Mosher, L., Moulton, S., Murari, D., Muraro, A., Murphy, A., Asakura, S., N. N., Na, Nabais, Y. S., Naish, F., Nakano, R., Nardon, T., Naulin, E., Nave, V., Nedzelski, M. F. F., Nemtsev, I., Nespoli, G., Neto, F., Neu, A., Neverov, R., Newman, V. S., Nicholls, M., Nicolas, K. J., Nielsen, T., Nielsen, A. H., Nilsson, P., Nishijima, E., Noble, D., Nocente, C., Nodwell, M., Nordlund, D., Nordman, K., Nouailletas, H., Nunes, R., Oberkofler, I., Odupitan, M., Ogawa, T., O'Gorman, M. T., Okabayashi, T., Olney, M., Omolayo, R., O'Mullane, O., Ongena, M., Orsitto, J., Orszagh, F., Oswuigwe, J., Otin, B. I., Owen, R., Paccagnella, A., Pace, R., Pacella, N., Packer, D., Page, L. W., Pajuste, A., Palazzo, E., Pamela, S., Panja, S., Papp, S., Paprok, P., Parail, R., Park, V., Parra, Diaz, Parsons, F., Pasqualotto, M., Patel, R., Pathak, A., Paton, S., Patten, D., Pau, H., Pawelec, A., Paz, Soldan, Peackoc, C., Pearson, A., Pehkonen, I. J., Peluso, S. P., Penot, E., Pereira, C., Pereira, A., Pereira, Puglia, P. P., Perez Von Thun, Peruzzo, C., Peschanyi, S., Peterka, S., Petersson, M., Petravich, P., Petre, G., Petrella, A., Petrå¾ilka, N., Peysson, V., Pfefferlã©, Y., Philipps, D., Pillon, V., Pintsuk, M., Piovesan, G., Pires Dos Reis, Piron, Lidia, Pironti, A., Pisano, F., Pitts, R., Pizzo, F., Plyusnin, V., Pomaro, N., Pompilian, O. G., Pool, P. J., Popovichev, S., Porfiri, M. T., Porosnicu, C., Porton, M., Possnert, G., Potzel, S., Powell, T., Pozzi, J., Prajapati, V., Prakash, R., Prestopino, G., Price, D., Price, M., Price, R., Prior, P., Proudfoot, R., Pucella, G., Puglia, P., Puiatti, M. E., Pulley, D., Purahoo, K., Pã¼tterich, T. h., Rachlew, E., Rack, M., Ragona, R., Rainford, M. S. J., Rakha, A., Ramogida, G., Ranjan, S., Rapson, C. J., Rasmussen, J. J., Rathod, K., Rattã¡, G., Ratynskaia, S., Ravera, G., Rayner, C., Rebai, M., Reece, D., Reed, A., Rã©fy, D., Regan, B., Regaã±a, J., Reich, M., Reid, N., Reimold, F., Reinhart, M., Reinke, M., Reiser, D., Rendell, D., Reux, C., Reyes, Cortes, Reynolds, S. D. A., Riccardo, S., Richardson, V., Riddle, N., Rigamonti, K., Rimini, D., Risner, F. G., Riva, J., Roach, M., Robins, C., Robinson, R. J., Robinson, S. A., Robson, T., Roccella, D. W., Rodionov, R., Rodrigues, R., Rodriguez, P., Rohde, J., Romanelli, V., Romanelli, F., Romanelli, M., Romazanov, S., Rowe, J., Rubel, S., Rubinacci, M., Rubino, G., Ruchko, G., Ruiz, L., Ruset, M., Rzadkiewicz, C., Saarelma, J., Sabot, S., Safi, R., Sagar, E., Saibene, P., Saint, Laurent, Salewski, F., Salmi, M., Salmon, A., Salzedas, R., Samaddar, F., Samm, D., Sandiford, U., Santa, D., Santala, P., Santos, M. I. K., Santucci, B., Sartori, A., Sartori, F., Sauter, R., Scannell, O., Schlummer, R., Schmid, T., Schmidt, K., Schmuck, V., Schneider, S., Schã¶pf, M., Schwã¶rer, K., Scott, D., Sergienko, S. D., Sertoli, G., Shabbir, M., Sharapov, A., Shaw, S. E., Shaw, A., Sheikh, R., Shepherd, H., Shevelev, A., Shumack, A., Sias, A., Sibbald, G., Sieglin, M., Silburn, B., Silva, S., Silva, A., Simmons, C., Simpson, P. A., Simpson, Hutchinson, Sinha, J., Sipilã¤, A., Sips, S. K., Sirã©n, A. C. C., Sirinelli, P., Sjã¶strand, A., Skiba, H., Skilton, M., Slabkowska, R., Slade, K., Smith, B., Smith, N., Smith, P. G., Smith, R., Smithies, T. J., Snoj, M., Soare, L., Solano, S., Somers, E. R., Sommariva, A., Sonato, C., Piergiorgio, Sopplesa, Sousa, A., Sozzi, J., Spagnolo, C., Silvia, Spelzini, Spineanu, T., Stables, F., Stamatelatos, G., Stamp, I., Staniec, M. F., Stankå«nas, P., Stan, Sion, Stead, C., Stefanikova, M. J., Stepanov, E., Stephen, I., Stephen, A. V., Stevens, M., Stevens, A., Strachan, B. D., Strand, J., Strauss, P., Strã¶m, H. R., Stubbs, P., Studholme, G., Subba, W., Summers, F., Svensson, H. P., Åšwiderski, J., Szabolics, Å. ., Szawlowski, T., Szepesi, M., Suzuki, G., Tã¡l, T. T., Tala, B., Talbot, T., Talebzadeh, A. R., Taliercio, S., Cesare, Tamain, Tame, P., Tang, C., Tardocchi, W., Taroni, M., Taylor, L., Taylor, D., Tegnered, K. A., Telesca, D., Teplova, G., Terranova, N., David, Testa, Tholerus, D., Thomas, E., Thomas, J., Thomas, J. D., Thompson, P., Thompson, A., Thompson, C. A., Thorne, V. K., Thornton, L., Thrysã¸e, A., Tigwell, A. S., Tipton, P. A., Tiseanu, N., Tojo, I., Tokitani, H., Tolias, M., Tomeå¡, P., Tonner, M., Towndrow, P., Trimble, M., Tripsky, P., Tsalas, M., Tsavalas, M., Tskhakaya, Jun, Turner, D., Turner, I., Turnyanskiy, M. M., Tvalashvili, M., Tyrrell, G., Uccello, S. G. J., Abidin, Ul, Uljanovs, Z., Ulyatt, J., Urano, D., Uytdenhouwen, H., Vadgama, I., Valcarcel, A. P., Valentinuzzi, D., Valisa, M., Vallejos, Olivares, Valovic, P., Van De Mortel, Van, Eester, Van, Renterghem, Van, Rooij, Varje, G. J., Varoutis, J., Vartanian, S., Vasava, S., Vasilopoulou, K., Vega, T., Verdoolaege, J., Verhoeven, G., Verona, R., Verona, Rinati, Veshchev, G., Vianello, E., Vicente, N., Viezzer, J., Villari, E., Villone, S., Vincenzi, F., Pietro, Vinyar, Viola, I., Vitins, B., Vizvary, A., Vlad, Z., Voitsekhovitch, M., Vondrã¡äek, I., Vora, P., Vu, N., Pires De Sa, Wakeling, W. W., Waldon, B., Walkden, C. W. F., Walker, N., Walker, M., Walsh, R., Wang, M., Wang, E., Warder, N., Warren, S., Waterhouse, R. J., Watkins, J., Watts, N. W., Wauters, C., Weckmann, T., Weiland, A., Weisen, J., Weiszflog, H., Wellstood, M., West, C., Wheatley, A. T., Whetham, M. R., Whitehead, S., Whitehead, A. M., Widdowson, B. D., Wiesen, A. M., Wilkinson, S., Williams, J., Wilson, M., Wilson, A. R., Wilson, D. J., Wilson, H. R., Wischmeier, J., Withenshaw, M., Withycombe, G., Witts, A., Wood, D. M., Wood, D., Woodley, R., Wray, C., Wright, S., Wright, J., J. C., Wu, Wukitch, J., Wynn, S., Xu, A., Yadikin, T., Yanling, D., Yao, W., Yavorskij, L., Yoo, V., Young, M. G., Young, C., Young, D., Young, I. D., Zacks, R., Zagorski, J., Zaitsev, R., Zanino, F. S., Zarins, R., Zastrow, A., Zerbini, K. D., Zhang, M., Zhou, W., Zilli, Y., Zoita, E., Zoletnik, V., Zychor, S., I, and JET Contributors
- Subjects
Jet (fluid) ,Surface analysis ,Materials science ,Divertor ,JET-ILW ,Material deposition ,Material erosion ,Nuclear engineering ,Condensed Matter Physics ,01 natural sciences ,Atomic and Molecular Physics, and Optics ,010305 fluids & plasmas ,13. Climate action ,Material Erosion ,0103 physical sciences ,Erosion ,010306 general physics ,Deposition (chemistry) ,Mathematical Physics - Abstract
Erosion of plasma-facing materials and successive transport and redeposition of eroded material are crucial processes determining the lifetime of plasma-facing components and the trapped tritium inventory in redeposited material layers. Erosion and deposition in the JET divertor were studied during the second JET ITER-like wall campaign ILW-2 in 2013-2014 by using a poloidal row of specially prepared divertor marker tiles including the tungsten bulk tile 5. The marker tiles were analyzed using elastic backscattering with 3-4.5 MeV incident protons and nuclear reaction analysis using 0.8-4.5 MeV 3He ions before and after the campaign. The erosion/deposition pattern observed during ILW-2 is qualitatively comparable to the first campaign ILW-1 in 2011-2012: deposits consist mainly of beryllium with 5-20 at.% of carbon and oxygen and small amounts of Ni and W. The highest deposition with deposited layer thicknesses up to 30 μm per campaign is still observed on the upper and horizontal parts of the inner divertor. Outer divertor tiles 5, 6, 7 and 8 are net W erosion areas. The observed D inventory is roughly comparable to the inventory observed during ILW-1. The results obtained during ILW-2 therefore confirm the positive results observed in ILW-1 with respect to reduced material deposition and hydrogen isotopes retention in the divertor.
- Published
- 2017
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5. Overview of fuel inventory in JET with the ITER-like wall
- Author
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Widdowson, A., Coad, J. P., Alves, E., Baron-Wiechec, A., Barradas, N. P., Brezinsek, S., Catarino, N., Corregidor, V., Heinola, K., Koivuranta, S., Krat, S., Lahtinen, A., Likonen, J., Matthews, G. F., Mayer, M., Petersson, P., Litaudon, Rubel M., Abduallev, X., Abhangi, S., Abreu, M., Afzal, P., Aggarwal, M., Ahlgren, K. M., Ahn, T., J. H., Aho, Mantila, Aiba, L., Airila, N., Albanese, M., Aldred, R., Alegre, V., Alessi, D., Aleynikov, E., Alfier, P., Alberto, Alkseev, Allinson, A., Alper, M., Alves, B., Ambrosino, E., Ambrosino, G., Amicucci, R., Amosov, L., Andersson, Sundã©n, Angelone, E., Anghel, M., Angioni, M., Appel, C., Appelbee, L., Arena, C., Ariola, P., Arnichand, M., Arshad, H., Ash, S., Ashikawa, A., Aslanyan, N., Asunta, V., Auriemma, O., Fulvio, Austin, Avotina, Y., Axton, L., Ayres, M. D., Bacharis, C., Baciero, M., Baiã¡o, A., Bailey, D., Baker, S., Balboa, A., Balden, I., Balshaw, M., Bament, N., Banks, R., Baranov, J. W., Barnard, Y. F., Barnes, M. 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R., Meshchaninov, P. h., Messiaen, S., Meyer, A., Mianowski, H., Michling, S., Middleton, Gear, Miettunen, D., Militello, J., Militello, Asp, Miloshevsky, E., Mink, G., Minucci, F., Miyoshi, S., Mlynã¡å™, Y., Molina, J., Monakhov, D., Moneti, I., Mooney, M., Moradi, R., Mordijck, S., Moreira, S., Moreno, L., Moro, R., Morris, F., Morris, A. W., Moser, J., Mosher, L., Moulton, S., Murari, D., Muraro, A., Murphy, A., Asakura, S., N. N., Na, Nabais, Y. S., Naish, F., Nakano, R., Nardon, T., Naulin, E., Nave, V., Nedzelski, M. F. F., Nemtsev, I., Nespoli, G., Neto, F., Neu, A., Neverov, R., Newman, V. S., Nicholls, M., Nicolas, K. J., Nielsen, T., Nielsen, A. H., Nilsson, P., Nishijima, E., Noble, D., Nocente, C., Nodwell, M., Nordlund, D., Nordman, K., Nouailletas, H., Nunes, R., Oberkofler, I., Odupitan, M., Ogawa, T., O'Gorman, M. T., Okabayashi, T., Olney, M., Omolayo, R., O'Mullane, O., Ongena, M., Orsitto, J., Orszagh, F., Oswuigwe, J., Otin, B. 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- Subjects
Nuclear and High Energy Physics ,Jet (fluid) ,Hydrogen ,Plasma parameters ,JET ITER-like wall ,Divertor ,Nuclear engineering ,chemistry.chemical_element ,Condensed Matter Physics ,01 natural sciences ,fuel retention ,010305 fluids & plasmas ,material migration ,chemistry ,Sputtering ,visual_art ,0103 physical sciences ,visual_art.visual_art_medium ,Environmental science ,Tile ,010306 general physics - Abstract
Post mortem analyses of JET ITER-Like-Wall tiles and passive diagnostics have been completed after each of the first two campaigns (ILW-1 and ILW-2). They show that the global fuel inventory is still dominated by co-deposition; hence plasma parameters and sputtering processes affecting material migration influence the distribution of retained fuel. In particular, differences between results from the two campaigns may be attributed to a greater proportion of pulses run with strike points in the divertor corners, and having about 300 discharges in hydrogen at the end of ILW-2. Recessed and remote areas can contribute to fuel retention due to the larger areas involved, e.g. recessed main chamber walls, gaps in castellated Be main chamber tiles and material migration to remote divertor areas. The fuel retention and material migration due to the bulk W Tile 5 during ILW-1 are presented. Overall these tiles account for only a small percentage of the global accountancy for ILW-1.
- Published
- 2017
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6. MeV-range velocity-space tomography from gamma-ray and neutron emission spectrometry measurements at JET
- Author
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Salewski, M., Nocente, M., Jacobsen, A. S., Binda, F., Cazzaniga, C., Ericsson, G., Eriksson, J., Gorini, G., Hellesen, C., Hjalmarsson, A., Kiptily, V. G., Koskela, T., Korsholm, S. B., Kurki-Suonio, T., Leipold, F., Madsen, J., Moseev, D., Nielsen, S. K., Rasmussen, J., Schneider, M., Sharapov, S. E., Stejner, M., Litaudon, Tardocchi M., Abduallev, X., Abhangi, S., Abreu, M., Afzal, P., Aggarwal, M., Ahlgren, K. M., Ahn, T., J. H., Aho, Mantila, Aiba, L., Airila, N., Albanese, M., Aldred, R., Alegre, V., Alessi, D., Aleynikov, E., Alfier, P., Alberto, Alkseev, Allinson, A., Alper, M., Alves, B., Ambrosino, E., Ambrosino, G., Amicucci, R., Amosov, L., Andersson, Sundã©n, Angelone, E., Anghel, M., Angioni, M., Appel, C., Appelbee, L., Arena, C., Ariola, P., Arnichand, M., Arshad, H., Ash, S., Ashikawa, A., Aslanyan, N., Asunta, V., Auriemma, O., Fulvio, Austin, Avotina, Y., Axton, L., Ayres, M. 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Nuclear and High Energy Physics ,gamma-ray spectrometry ,Neutron emission ,Fluids & Plasmas ,Astrophysics::High Energy Astrophysical Phenomena ,Nuclear Theory ,01 natural sciences ,7. Clean energy ,Atomic ,010305 fluids & plasmas ,Ion ,Nuclear physics ,Particle and Plasma Physics ,Physics::Plasma Physics ,0103 physical sciences ,fast ion ,γ-ray spectrometry ,Neutron ,Nuclear ,Emission spectrum ,fast ions ,010306 general physics ,Nuclear Experiment ,tokamak ,Nuclear and High Energy Physic ,Physics ,Jet (fluid) ,Neutron stimulated emission computed tomography ,Gamma ray ,Molecular ,neutron emission spectrometry ,velocity-space tomography ,Condensed Matter Physics ,Physics::Accelerator Physics ,Atomic physics ,Ion cyclotron resonance - Abstract
© 2017 Technical University of Denmark. We demonstrate the measurement of a 2D MeV-range ion velocity distribution function by velocity-space tomography at JET. Deuterium ions were accelerated into the MeV-range by third harmonic ion cyclotron resonance heating. We made measurements with three neutron emission spectrometers and a high-resolution γ-ray spectrometer detecting the γ-rays released in two reactions. The tomographic inversion based on these five spectra is in excellent agreement with numerical simulations with the ASCOT-RFOF and the SPOT-RFOF codes. The length of the measured fast-ion tail corroborates the prediction that very few particles are accelerated above 2 MeV due to the weak wave-particle interaction at higher energies.
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- 2017
- Full Text
- View/download PDF
7. Ph1 affects remodelling of the heterochromatin at the telomeres and the centromeres at the onset of meiosis
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Prieto P, Colas I, Shaw P, and Moore G
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Wheat breeding ,Wheat genetics - Published
- 2008
8. Mass spectrometry in plant proteomic analysis
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Colas, I., primary, Koroleva, O., additional, and Shaw, P. J., additional
- Published
- 2010
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9. Image analysis workflows to reveal the spatial organization of cell nuclei and chromosomes.
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Randall RS, Jourdain C, Nowicka A, Kaduchová K, Kubová M, Ayoub MA, Schubert V, Tatout C, Colas I, Kalyanikrishna, Desset S, Mermet S, Boulaflous-Stevens A, Kubalová I, Mandáková T, Heckmann S, Lysak MA, Panatta M, Santoro R, Schubert D, Pecinka A, Routh D, and Baroux C
- Subjects
- Humans, Workflow, In Situ Hybridization, Fluorescence, Microscopy, Fluorescence, Green Fluorescent Proteins, Cell Nucleus, Chromatin
- Abstract
Nucleus, chromatin, and chromosome organization studies heavily rely on fluorescence microscopy imaging to elucidate the distribution and abundance of structural and regulatory components. Three-dimensional (3D) image stacks are a source of quantitative data on signal intensity level and distribution and on the type and shape of distribution patterns in space. Their analysis can lead to novel insights that are otherwise missed in qualitative-only analyses. Quantitative image analysis requires specific software and workflows for image rendering, processing, segmentation, setting measurement points and reference frames and exporting target data before further numerical processing and plotting. These tasks often call for the development of customized computational scripts and require an expertise that is not broadly available to the community of experimental biologists. Yet, the increasing accessibility of high- and super-resolution imaging methods fuels the demand for user-friendly image analysis workflows. Here, we provide a compendium of strategies developed by participants of a training school from the COST action INDEPTH to analyze the spatial distribution of nuclear and chromosomal signals from 3D image stacks, acquired by diffraction-limited confocal microscopy and super-resolution microscopy methods (SIM and STED). While the examples make use of one specific commercial software package, the workflows can easily be adapted to concurrent commercial and open-source software. The aim is to encourage biologists lacking custom-script-based expertise to venture into quantitative image analysis and to better exploit the discovery potential of their images. Abbreviations: 3D FISH: three-dimensional fluorescence in situ hybridization; 3D: three-dimensional; ASY1: ASYNAPTIC 1; CC: chromocenters; CO: Crossover; DAPI: 4',6-diamidino-2-phenylindole; DMC1: DNA MEIOTIC RECOMBINASE 1; DSB: Double-Strand Break; FISH: fluorescence in situ hybridization; GFP: GREEN FLUORESCENT PROTEIN; HEI10: HUMAN ENHANCER OF INVASION 10; NCO: Non-Crossover; NE: Nuclear Envelope; Oligo-FISH: oligonucleotide fluorescence in situ hybridization; RNPII: RNA Polymerase II; SC: Synaptonemal Complex; SIM: structured illumination microscopy; ZMM (ZIP: MSH4: MSH5 and MER3 proteins); ZYP1: ZIPPER-LIKE PROTEIN 1.
- Published
- 2022
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10. Location and Identification on Chromosome 3B of Bread Wheat of Genes Affecting Chiasma Number.
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Darrier B, Colas I, Rimbert H, Choulet F, Bazile J, Sortais A, Jenczewski E, and Sourdille P
- Abstract
Understanding meiotic crossover (CO) variation in crops like bread wheat ( Triticum aestivum L.) is necessary as COs are essential to create new, original and powerful combinations of genes for traits of agronomical interest. We cytogenetically characterized a set of wheat aneuploid lines missing part or all of chromosome 3B to identify the most influential regions for chiasma formation located on this chromosome. We showed that deletion of the short arm did not change the total number of chiasmata genome-wide, whereas this latter was reduced by ~35% while deleting the long arm. Contrary to what was hypothesized in a previous study, deletion of the long arm does not disturb the initiation of the synaptonemal complex (SC) in early meiotic stages. However, progression of the SC is abnormal, and we never observed its completion when the long arm is deleted. By studying six different deletion lines (missing different parts of the long arm), we revealed that at least two genes located in both the proximal (C-3BL2-0.22) and distal (3BL7-0.63-1.00) deletion bins are involved in the control of chiasmata, each deletion reducing the number of chiasmata by ~15%. We combined sequence analyses of deletion bins with RNA-Seq data derived from meiotic tissues and identified a set of genes for which at least the homoeologous copy on chromosome 3B is expressed and which are involved in DNA processing. Among these genes, eight (CAP-E1/E2, DUO1, MLH1, MPK4, MUS81, RTEL1, SYN4, ZIP4) are known to be involved in the recombination pathway., Competing Interests: The authors declare no conflict of interest.
- Published
- 2022
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11. The proteome of developing barley anthers during meiotic prophase I.
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Lewandowska D, Orr J, Schreiber M, Colas I, Ramsay L, Zhang R, and Waugh R
- Subjects
- Flowers, Meiosis, Meiotic Prophase I, Plant Proteins genetics, Plant Proteins metabolism, Hordeum genetics, Hordeum metabolism, Proteome metabolism
- Abstract
Flowering plants reproduce sexually by combining a haploid male and female gametophyte during fertilization. Male gametophytes are localized in the anthers, each containing reproductive (meiocyte) and non-reproductive tissue necessary for anther development and maturation. Meiosis, where chromosomes pair and exchange their genetic material during a process called recombination, is one of the most important and sensitive stages in breeding, ensuring genetic diversity. Most anther development studies have focused on transcript variation, but very few have been correlated with protein abundance. Taking advantage of a recently published barley anther transcriptomic (BAnTr) dataset and a newly developed sensitive mass spectrometry-based approach to analyse the barley anther proteome, we conducted high-resolution mass spectrometry analysis of barley anthers, collected at six time points and representing their development from pre-meiosis to metaphase. Each time point was carefully staged using immunocytology, providing a robust and accurate staging mirroring our previous BAnTr dataset. We identified >6100 non-redundant proteins including 82 known and putative meiotic proteins. Although the protein abundance was relatively stable throughout prophase I, we were able to quantify the dynamic variation of 336 proteins. We present the first quantitative comparative proteomics study of barley anther development during meiotic prophase I when the important process of homologous recombination is taking place., (© The Author(s) 2021. Published by Oxford University Press on behalf of the Society for Experimental Biology.)
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- 2022
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12. An Induced Mutation in HvRECQL4 Increases the Overall Recombination and Restores Fertility in a Barley HvMLH3 Mutant Background.
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Arrieta M, Macaulay M, Colas I, Schreiber M, Shaw PD, Waugh R, and Ramsay L
- Abstract
Plant breeding relies on the meiotic recombination or crossing over to generate the new combinations of the alleles along and among the chromosomes. However, crossing over is constrained in the crops such as barley by a combination of the low frequency and biased distribution. In this study, we attempted to identify the genes that limit the recombination by performing a suppressor screen for the restoration of fertility to the semi-fertile barley mutant desynaptic10 ( des10 ), carrying a mutation in the barley ortholog of MutL-Homolog 3 ( HvMLH3 ), a member of the MutL-homolog (MLH) family of DNA mismatch repair genes. des10 mutants exhibit reduced recombination and fewer chiasmata, resulting in the loss of obligate crossovers (COs) leading to chromosome mis-segregation. We identified several candidate suppressor lines and confirmed their restored fertility in an Hvmlh3 background in the subsequent generations. We focus on one of the candidate suppressor lines, SuppLine2099 , which showed the most complete restoration of fertility. We characterized this line by using a target-sequence enrichment and sequencing (TENSEQ) capture array representing barley orthologs of 46 meiotic genes. We found that SuppLine2099 contained a C/T change in the anti-CO gene RecQ-like helicase 4 (RECQL4) resulting in the substitution of a non-polar glycine to a polar aspartic acid (G700D) amino acid in the conserved helicase domain. Single nucleotide polymorphism (SNP) genotyping of F
3 populations revealed a significant increase in the recombination frequency in lines with Hvrecql4 in the Hvmlh3 background that was associated with the restoration of fertility. The genotyping also indicated that there was nearly double the recombination levels in homozygous Hvrecql4 lines compared to the wild type (WT). However, we did not observe any significant change in the distribution of CO events. Our results confirm the anti-CO role of RECQL4 in a large genome cereal and establish the possibility of testing the utility of increasing recombination in the context of traditional crop improvement., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Arrieta, Macaulay, Colas, Schreiber, Shaw, Waugh and Ramsay.)- Published
- 2021
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13. Ubiquitination in Plant Meiosis: Recent Advances and High Throughput Methods.
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Orr JN, Waugh R, and Colas I
- Abstract
Meiosis is a specialized cell division which is essential to sexual reproduction. The success of this highly ordered process involves the timely activation, interaction, movement, and removal of many proteins. Ubiquitination is an extraordinarily diverse post-translational modification with a regulatory role in almost all cellular processes. During meiosis, ubiquitin localizes to chromatin and the expression of genes related to ubiquitination appears to be enhanced. This may be due to extensive protein turnover mediated by proteasomal degradation. However, degradation is not the only substrate fate conferred by ubiquitination which may also mediate, for example, the activation of key transcription factors. In plant meiosis, the specific roles of several components of the ubiquitination cascade-particularly SCF complex proteins, the APC/C, and HEI10-have been partially characterized indicating diverse roles in chromosome segregation, recombination, and synapsis. Nonetheless, these components remain comparatively poorly understood to their counterparts in other processes and in other eukaryotes. In this review, we present an overview of our understanding of the role of ubiquitination in plant meiosis, highlighting recent advances, remaining challenges, and high throughput methods which may be used to overcome them., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Orr, Waugh and Colas.)
- Published
- 2021
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14. Barley Anther and Meiocyte Transcriptome Dynamics in Meiotic Prophase I.
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Barakate A, Orr J, Schreiber M, Colas I, Lewandowska D, McCallum N, Macaulay M, Morris J, Arrieta M, Hedley PE, Ramsay L, and Waugh R
- Abstract
In flowering plants, successful germinal cell development and meiotic recombination depend upon a combination of environmental and genetic factors. To gain insights into this specialized reproductive development program we used short- and long-read RNA-sequencing (RNA-seq) to study the temporal dynamics of transcript abundance in immuno-cytologically staged barley ( Hordeum vulgare ) anthers and meiocytes. We show that the most significant transcriptional changes in anthers occur at the transition from pre-meiosis to leptotene-zygotene, which is followed by increasingly stable transcript abundance throughout prophase I into metaphase I-tetrad. Our analysis reveals that the pre-meiotic anthers are enriched in long non-coding RNAs (lncRNAs) and that entry to meiosis is characterized by their robust and significant down regulation. Intriguingly, only 24% of a collection of putative meiotic gene orthologs showed differential transcript abundance in at least one stage or tissue comparison. Argonautes, E3 ubiquitin ligases, and lys48 specific de-ubiquitinating enzymes were enriched in prophase I meiocyte samples. These developmental, time-resolved transcriptomes demonstrate remarkable stability in transcript abundance in meiocytes throughout prophase I after the initial and substantial reprogramming at meiosis entry and the complexity of the regulatory networks involved in early meiotic processes., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Barakate, Orr, Schreiber, Colas, Lewandowska, McCallum, Macaulay, Morris, Arrieta, Hedley, Ramsay and Waugh.)
- Published
- 2021
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15. The effect of heat stress on sugar beet recombination.
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Arrieta M, Willems G, DePessemier J, Colas I, Burkholz A, Darracq A, Vanstraelen S, Pacolet P, Barré C, Kempeneers P, Waugh R, Barnes S, and Ramsay L
- Subjects
- Beta vulgaris physiology, Genotype, Meiosis, Plant Breeding, Beta vulgaris genetics, Crossing Over, Genetic, Hot Temperature, Stress, Physiological
- Abstract
Meiotic recombination plays a crucial role in plant breeding through the creation of new allelic combinations. Therefore, lack of recombination in some genomic regions constitutes a constraint for breeding programmes. In sugar beet, one of the major crops in Europe, recombination occurs mainly in the distal portions of the chromosomes, and so the development of simple approaches to change this pattern is of considerable interest for future breeding and genetics. In the present study, the effect of heat stress on recombination in sugar beet was studied by treating F
1 plants at 28 °C/25 °C (day/night) and genotyping the progeny. F1 plants were reciprocally backcrossed allowing the study of male and female meiosis separately. Genotypic data indicated an overall increase in crossover frequency of approximately one extra crossover per meiosis, with an associated increase in pericentromeric recombination under heat treatment. Our data indicate that the changes were mainly induced by alterations in female meiosis only, showing that heterochiasmy in sugar beet is reduced under heat stress. Overall, despite the associated decrease in fertility, these data support the potential use of heat stress to foster recombination in sugar beet breeding programmes.- Published
- 2021
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16. Chromatin dynamics during interphase and cell division: similarities and differences between model and crop plants.
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Pecinka A, Chevalier C, Colas I, Kalantidis K, Varotto S, Krugman T, Michailidis C, Vallés MP, Muñoz A, and Pradillo M
- Subjects
- Cell Division, Chromosomes, Interphase, Chromatin genetics, Plant Breeding
- Abstract
Genetic information in the cell nucleus controls organismal development and responses to the environment, and finally ensures its own transmission to the next generations. To achieve so many different tasks, the genetic information is associated with structural and regulatory proteins, which orchestrate nuclear functions in time and space. Furthermore, plant life strategies require chromatin plasticity to allow a rapid adaptation to abiotic and biotic stresses. Here, we summarize current knowledge on the organization of plant chromatin and dynamics of chromosomes during interphase and mitotic and meiotic cell divisions for model and crop plants differing as to genome size, ploidy, and amount of genomic resources available. The existing data indicate that chromatin changes accompany most (if not all) cellular processes and that there are both shared and unique themes in the chromatin structure and global chromosome dynamics among species. Ongoing efforts to understand the molecular mechanisms involved in chromatin organization and remodeling have, together with the latest genome editing tools, potential to unlock crop genomes for innovative breeding strategies and improvements of various traits., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Experimental Biology.)
- Published
- 2020
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17. A Modular Tray Growth System for Barley.
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Arrieta M, Colas I, Macaulay M, Waugh R, and Ramsay L
- Subjects
- Chromosomes, Plant, Immunohistochemistry, Phenotype, Plant Development genetics, Recombination, Genetic, Stress, Physiological, Temperature, Hordeum genetics, Hordeum growth & development, Meiosis genetics
- Abstract
Determining when a barley plant starts and finishes meiosis is not trivial as when the spikelets undergo meiosis, the spike is not visible as it is still well within the leaf sheath on the developing tiller. This is a general constraint for any experiment involving meiosis, such as cytology, RNA extractions, or abiotic stress treatments aiming to target such a developmental stage. The lack of synchronicity between barley tillers within the same plant exacerbates the difficulty to determine the overall meiotic stage of a plant at a certain time.Given the lack of a nondestructive staging system for predicting the entry into meiosis and the problems of working with large pot plant systems, a modular plant growing is proposed. This system enables the growth of a high number of plants in a small surface, each producing a single tiller. The modular tray system was used to generate a nondestructive prediction tool for meiosis by using external morphological features. As an example, the system is used here for heat treating F
1 plants in early meiosis stages to modify recombination.- Published
- 2020
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18. Following the Formation of Synaptonemal Complex Formation in Wheat and Barley by High-Resolution Microscopy.
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Darrier B, Arrieta M, Mittmann SU, Sourdille P, Ramsay L, Waugh R, and Colas I
- Subjects
- Fluorescent Antibody Technique methods, Imaging, Three-Dimensional, Chromosome Pairing, Hordeum genetics, Meiosis, Microscopy methods, Synaptonemal Complex, Triticum genetics
- Abstract
Wheat and barley have large genomes of 15 Gb and 5.1 Gb, respectively, which is much larger than the human genome (3.3 Gb). The release of their respective genomes has been a tremendous advance the understanding of the genome organization and the ability for deeper functional analysis in particular meiosis. Meiosis is the cell division required during sexual reproduction. One major event of meiosis is called recombination, or the formation of crossing over, a tight link between homologous chromosomes, ensuring gene exchange and faithful chromosome segregation. Recombination is a major driver of genetic diversity but in these large genome crops, the vast majority of these events is constrained at the end of their chromosomes. It is estimated that in barley, about 30% of the genes are located within the poor recombining centromeric regions, making important traits, such as resistance to pest and disease for example, difficult to access. Increasing recombination in these crops has the potential to speed up breeding program and requires a good understand of the meiotic mechanism. However, most research on recombination in plant has been carried in Arabidopsis thaliana which despite many of the advantages it brings for plant research, has a small genome and more spread out of recombination compare to barley or wheat. Advance in microscopy and cytological procedures have emerged in the last few years, allowing to follow meiotic events in these crops. This protocol provides the steps required for cytological preparation of barley and wheat pollen mother cells for light microscopy, highlighting some of the differences between the two cereals.
- Published
- 2020
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19. desynaptic5 carries a spontaneous semi-dominant mutation affecting Disrupted Meiotic cDNA 1 in barley.
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Colas I, Barakate A, Macaulay M, Schreiber M, Stephens J, Vivera S, Halpin C, Waugh R, and Ramsay L
- Subjects
- Amino Acid Sequence, Base Sequence, Hordeum metabolism, Mutation genetics, Phylogeny, Plant Proteins chemistry, Plant Proteins metabolism, Sequence Alignment, Hordeum genetics, Plant Proteins genetics
- Abstract
Despite conservation of the process of meiosis, recombination landscapes vary between species, with large genome grasses such as barley (Hordeum vulgare L.) exhibiting a pattern of recombination that is very heavily skewed to the ends of chromosomes. We have been using a collection of semi-sterile desynaptic meiotic mutant lines to help elucidate how recombination is controlled in barley and the role of the corresponding wild-type (WT) meiotic genes within this process. Here we applied a combination of genetic segregation analysis, cytogenetics, and immunocytology to genetically map and characterize the meiotic mutant desynaptic5 (des5). We identified an exonic insertion in the positional candidate ortholog of Disrupted Meiotic cDNA 1 (HvDMC1) on chromosome 5H of des5. des5 exhibits a severe meiotic phenotype with disturbed synapsis, reduced crossovers, and chromosome mis-segregation. The meiotic phenotype and reduced fertility of des5 is similarly observed in Hvdmc1RNAi transgenic plants and HvDMC1p:GusPlus reporter lines show DMC1 expression specifically in the developing inflorescence. The des5 mutation maintains the reading frame of the gene and exhibits semi-dominance with respect to recombination in the heterozygote indicating the value of non-knockout mutations for dissection of the control of recombination in the early stages of meiosis., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Experimental Biology.)
- Published
- 2019
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20. Application of a Sensitive and Reproducible Label-Free Proteomic Approach to Explore the Proteome of Individual Meiotic-Phase Barley Anthers.
- Author
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Lewandowska D, Zhang R, Colas I, Uzrek N, and Waugh R
- Abstract
Meiosis is a highly dynamic and precisely regulated process of cell division, leading to the production of haploid gametes from one diploid parental cell. In the crop plant barley ( Hordeum vulgare ), male meiosis occurs in anthers, in specialized cells called meiocytes. Barley meiotic tissue is scarce and not easily accessible, making meiosis study a challenging task. We describe here a new micro-proteomics workflow that allows sensitive and reproducible genome-wide label-free proteomic analysis of individual staged barley anthers. This micro-proteomic approach detects more than 4,000 proteins from such small amounts of material as two individual anthers, covering a dynamic range of protein relative abundance levels across five orders of magnitude. We applied our micro-proteomics workflow to investigate the proteome of the developing barley anther containing pollen mother cells in the early stages of meiosis and we successfully identified 57 known and putative meiosis-related proteins. Meiotic proteins identified in our study were found to be key players of many steps and processes in early prophase such as: chromosome condensation, synapsis, DNA double-strand breaks or crossover formation. Considering the small amount of starting material, this work demonstrates an important technological advance in plant proteomics and can be applied for proteomic examination of many size-limited plant specimens. Moreover, it is the first insight into the proteome of individual barley anther at early meiosis. The proteomic data have been deposited to the ProteomeXchange with the accession number PXD010887.
- Published
- 2019
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21. Preparation of Barley Pollen Mother Cells for Confocal and Super Resolution Microscopy.
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Mittmann S, Arrieta M, Ramsay L, Waugh R, and Colas I
- Subjects
- Chromosome Pairing, Meiosis, Hordeum cytology, Microscopy methods, Pollen cytology
- Abstract
Recombination (crossover) drives the release of genetic diversity in plant breeding programs. However, in barley, recombination is skewed toward the telomeric ends of its seven chromosomes, restricting the re-assortment of about 30% of the genes located in the centromeric regions of its large 5.1 Gb genome. A better understanding of meiosis and recombination could provide ways of modulating crossover distribution and frequency in barley as well as in other grasses, including wheat. While most research on recombination has been carried out in the model plant Arabidopsis thaliana, recent studies in barley (Hordeum Vulgare) have provided new insights into the control of crossing over in large genome species. A major achievement in these studies has been the use of cytological procedures to follow meiotic events. This protocol provides detailed practical steps required to perform immunostaining of barley meiocytes (pollen mother cells) for confocal or structured illumination microscopy.
- Published
- 2019
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22. Observation of Extensive Chromosome Axis Remodeling during the "Diffuse-Phase" of Meiosis in Large Genome Cereals.
- Author
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Colas I, Darrier B, Arrieta M, Mittmann SU, Ramsay L, Sourdille P, and Waugh R
- Abstract
The production of balanced fertile haploid gametes requires the faithful separation of paired (synapsed) chromosomes toward the end of meiotic prophase I (desynapsis). This involves the timely dissolution of the synaptonemal complex during the pachytene-diplotene transition, a stage traditionally referred to as the "diffuse stage." In species with large genomes such as, barley ( Hordeum vulgare L.) and wheat ( Triticum aestivum L.) we know most about the early stages of meiotic prophase I. There, synapsis initiates at the telomeric ends of chromosomes and progresses toward the centromeric regions through the ordered assembly of the synaptonemal complex (SC). Synapsis is impacted by recombination (crossing over, CO) which locally modifies the extent of chromatin compaction and extension. CO is uneven along the chromosomes, occurring mainly toward the telomeric regions resulting in a highly skewed distribution of recombination events. However, we know very little about the process of desynapsis which occurs during the "diffuse stage," where the synapsed and recombined chromosomes faithfully desynapse and separate into daughter cells. Here, using 3D-SIM super-resolution immuno-cytology combined with the use of antibodies directed against two crucial SC proteins, ASY1 and ZYP1, we followed the whole of meiosis I (i.e., both synapsis and desynapsis) in both barley and wheat. We showed that synapsis forms a characteristic tri-partite SC structure in zygotene (more clearly seen in barley). Toward the end of meiosis I, as the SC starts to disassemble, we show that extensive chromosome axis remodeling results in the formation of characteristic "tinsel-like" structures in both wheat and barley. By using a mutant ( des10 ) that is severely compromised in polymerization of ZYP1during synapsis, we show that tinsel structure formation during SC dissolution is not dependant on full synapsis and may relate instead to changes in expansion stress. Our observations highlight a potentially new role for ASYNAPSIS1 (ASY1) in desynapsis, in addition to chromosome synapsis and cohesion.
- Published
- 2017
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23. A spontaneous mutation in MutL-Homolog 3 (HvMLH3) affects synapsis and crossover resolution in the barley desynaptic mutant des10.
- Author
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Colas I, Macaulay M, Higgins JD, Phillips D, Barakate A, Posch M, Armstrong SJ, Franklin FC, Halpin C, Waugh R, and Ramsay L
- Subjects
- Base Sequence, Chromosome Mapping, Chromosome Segregation genetics, Chromosomes, Plant genetics, Crosses, Genetic, DNA Mismatch Repair genetics, Genes, Plant, Homologous Recombination genetics, Plant Proteins chemistry, Plant Proteins metabolism, Chromosome Pairing genetics, Crossing Over, Genetic, Hordeum genetics, Mutation genetics, Plant Proteins genetics
- Abstract
Although meiosis is evolutionarily conserved, many of the underlying mechanisms show species-specific differences. These are poorly understood in large genome plant species such as barley (Hordeum vulgare) where meiotic recombination is very heavily skewed to the ends of chromosomes. The characterization of mutant lines can help elucidate how recombination is controlled. We used a combination of genetic segregation analysis, cytogenetics, immunocytology and 3D imaging to genetically map and characterize the barley meiotic mutant DESYNAPTIC 10 (des10). We identified a spontaneous exonic deletion in the orthologue of MutL-Homolog 3 (HvMlh3) as the causal lesion. Compared with wild-type, des10 mutants exhibit reduced recombination and fewer chiasmata, resulting in the loss of obligate crossovers and leading to chromosome mis-segregation. Using 3D structured illumination microscopy (3D-SIM), we observed that normal synapsis progression was also disrupted in des10, a phenotype that was not evident with standard confocal microscopy and that has not been reported with Mlh3 knockout mutants in Arabidopsis. Our data provide new insights on the interplay between synapsis and recombination in barley and highlight the need for detailed studies of meiosis in nonmodel species. This study also confirms the importance of early stages of prophase I for the control of recombination in large genome cereals., (© 2016 The Authors. New Phytologist © 2016 New Phytologist Trust.)
- Published
- 2016
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24. Chromatin state analysis of the barley epigenome reveals a higher-order structure defined by H3K27me1 and H3K27me3 abundance.
- Author
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Baker K, Dhillon T, Colas I, Cook N, Milne I, Milne L, Bayer M, and Flavell AJ
- Subjects
- Chromatin genetics, Chromatin Immunoprecipitation, Epigenesis, Genetic genetics, Heterochromatin genetics, Histones genetics, Hordeum genetics, Chromatin metabolism, Heterochromatin metabolism, Histones metabolism, Hordeum metabolism
- Abstract
Combinations of histones carrying different covalent modifications are a major component of epigenetic variation. We have mapped nine modified histones in the barley seedling epigenome by chromatin immunoprecipitation next-generation sequencing (ChIP-seq). The chromosomal distributions of the modifications group them into four different classes, and members of a given class also tend to coincide at the local DNA level, suggesting that global distribution patterns reflect local epigenetic environments. We used this peak sharing to define 10 chromatin states representing local epigenetic environments in the barley genome. Five states map mainly to genes and five to intergenic regions. Two genic states involving H3K36me3 are preferentially associated with constitutive gene expression, while an H3K27me3-containing genic state is associated with differentially expressed genes. The 10 states display striking distribution patterns that divide barley chromosomes into three distinct global environments. First, telomere-proximal regions contain high densities of H3K27me3 covering both genes and intergenic DNA, together with very low levels of the repressive H3K27me1 modification. Flanking these are gene-rich interior regions that are rich in active chromatin states and have greatly decreased levels of H3K27me3 and increasing amounts of H3K27me1 and H3K9me2. Lastly, H3K27me3-depleted pericentromeric regions contain gene islands with active chromatin states separated by extensive retrotransposon-rich regions that are associated with abundant H3K27me1 and H3K9me2 modifications. We propose an epigenomic framework for barley whereby intergenic H3K27me3 specifies facultative heterochromatin in the telomere-proximal regions and H3K27me1 is diagnostic for constitutive heterochromatin elsewhere in the barley genome., (© 2015 The Authors The Plant Journal published by Society for Experimental Biology and John Wiley & Sons Ltd.)
- Published
- 2015
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25. The effect of temperature on the male and female recombination landscape of barley.
- Author
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Phillips D, Jenkins G, Macaulay M, Nibau C, Wnetrzak J, Fallding D, Colas I, Oakey H, Waugh R, and Ramsay L
- Subjects
- Cell Nucleus metabolism, Chromosome Mapping, Chromosomes, Plant genetics, Crosses, Genetic, Genetic Linkage, Genetic Loci, Hordeum cytology, Meiosis, Synaptonemal Complex, Hordeum genetics, Recombination, Genetic, Temperature
- Abstract
Barley (Hordeum vulgare) is a crop of global significance. However, a third of the genes of barley are largely inaccessible to conventional breeding programmes as crossovers are localised to the ends of the chromosomes. This work examines whether crossovers can be shifted to more proximal regions simply by elevating growth temperature. We utilised a genome-wide marker set for linkage analysis combined with cytological mapping of crossover events to examine the recombination landscape of plants grown at different temperatures. We found that barley shows heterochiasmy, that is, differences between female and male recombination frequencies. In addition, we found that elevated temperature significantly changes patterns of recombination in male meiosis only, with a repositioning of Class I crossovers determined by cytological mapping of HvMLH3 foci. We show that the length of synaptonemal complexes in male meiocytes increases in response to temperature. The results demonstrate that the distribution of crossover events are malleable and can be shifted to proximal regions by altering the growth temperature. The shift in recombination is the result of altering the distribution of Class I crossovers, but the higher recombination at elevated temperatures is potentially not the result of an increase in Class I events., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)
- Published
- 2015
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26. The low-recombining pericentromeric region of barley restricts gene diversity and evolution but not gene expression.
- Author
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Baker K, Bayer M, Cook N, Dreißig S, Dhillon T, Russell J, Hedley PE, Morris J, Ramsay L, Colas I, Waugh R, Steffenson B, Milne I, Stephen G, Marshall D, and Flavell AJ
- Subjects
- Base Sequence, Gene Duplication, Gene Expression, Gene Ontology, Heterochromatin genetics, Molecular Sequence Data, Recombination, Genetic, Sequence Analysis, RNA, Evolution, Molecular, Genetic Variation, Genome, Plant genetics, Hordeum genetics
- Abstract
The low-recombining pericentromeric region of the barley genome contains roughly a quarter of the genes of the species, embedded in low-recombining DNA that is rich in repeats and repressive chromatin signatures. We have investigated the effects of pericentromeric region residency upon the expression, diversity and evolution of these genes. We observe no significant difference in average transcript level or developmental RNA specificity between the barley pericentromeric region and the rest of the genome. In contrast, all of the evolutionary parameters studied here show evidence of compromised gene evolution in this region. First, genes within the pericentromeric region of wild barley show reduced diversity and significantly weakened purifying selection compared with the rest of the genome. Second, gene duplicates (ohnolog pairs) derived from the cereal whole-genome duplication event ca. 60MYa have been completely eliminated from the barley pericentromeric region. Third, local gene duplication in the pericentromeric region is reduced by 29% relative to the rest of the genome. Thus, the pericentromeric region of barley is a permissive environment for gene expression but has restricted gene evolution in a sizeable fraction of barley's genes., (© 2014 The Authors The Plant Journal © 2014 John Wiley & Sons Ltd.)
- Published
- 2014
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27. The synaptonemal complex protein ZYP1 is required for imposition of meiotic crossovers in barley.
- Author
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Barakate A, Higgins JD, Vivera S, Stephens J, Perry RM, Ramsay L, Colas I, Oakey H, Waugh R, Franklin FC, Armstrong SJ, and Halpin C
- Subjects
- Chromosomes, Plant genetics, DNA Breaks, Double-Stranded, Gene Expression Regulation, Plant, Gene Knockdown Techniques, Meiotic Prophase I, Molecular Sequence Data, Nondisjunction, Genetic, Phylogeny, Plant Proteins genetics, Plants, Genetically Modified, RNA Interference, RNA, Messenger genetics, RNA, Messenger metabolism, Time Factors, Crossing Over, Genetic, Hordeum cytology, Hordeum genetics, Meiosis genetics, Plant Proteins metabolism, Synaptonemal Complex metabolism
- Abstract
In many cereal crops, meiotic crossovers predominantly occur toward the ends of chromosomes and 30 to 50% of genes rarely recombine. This limits the exploitation of genetic variation by plant breeding. Previous reports demonstrate that chiasma frequency can be manipulated in plants by depletion of the synaptonemal complex protein ZIPPER1 (ZYP1) but conflict as to the direction of change, with fewer chiasmata reported in Arabidopsis thaliana and more crossovers reported for rice (Oryza sativa). Here, we use RNA interference (RNAi) to reduce the amount of ZYP1 in barley (Hordeum vulgare) to only 2 to 17% of normal zygotene levels. In the ZYP1(RNAi) lines, fewer than half of the chromosome pairs formed bivalents at metaphase and many univalents were observed, leading to chromosome nondisjunction and semisterility. The number of chiasmata per cell was reduced from 14 in control plants to three to four in the ZYP1-depleted lines, although the localization of residual chiasmata was not affected. DNA double-strand break formation appeared normal, but the recombination pathway was defective at later stages. A meiotic time course revealed a 12-h delay in prophase I progression to the first labeled tetrads. Barley ZYP1 appears to function similarly to ZIP1/ZYP1 in yeast and Arabidopsis, with an opposite effect on crossover number to ZEP1 in rice, another member of the Poaceae.
- Published
- 2014
- Full Text
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28. The Ph1 locus suppresses Cdk2-type activity during premeiosis and meiosis in wheat.
- Author
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Greer E, Martín AC, Pendle A, Colas I, Jones AM, Moore G, and Shaw P
- Subjects
- Cyclin-Dependent Kinase 2 genetics, Gene Expression Regulation, Plant genetics, Gene Expression Regulation, Plant physiology, Meiosis genetics, Molecular Sequence Data, Plant Proteins genetics, Triticum genetics, Cyclin-Dependent Kinase 2 metabolism, Meiosis physiology, Plant Proteins metabolism, Triticum metabolism
- Abstract
Despite possessing multiple sets of related (homoeologous) chromosomes, hexaploid wheat (Triticum aestivum) restricts pairing to just true homologs at meiosis. Deletion of a single major locus, Pairing homoeologous1 (Ph1), allows pairing of homoeologs. How can the same chromosomes be processed as homologs instead of being treated as nonhomologs? Ph1 was recently defined to a cluster of defective cyclin-dependent kinase (Cdk)-like genes showing some similarity to mammalian Cdk2. We reasoned that the cluster might suppress Cdk2-type activity and therefore affect replication and histone H1 phosphorylation. Our study does indeed reveal such effects, suggesting that Cdk2-type phosphorylation has a major role in determining chromosome specificity during meiosis.
- Published
- 2012
- Full Text
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29. Effective chromosome pairing requires chromatin remodeling at the onset of meiosis.
- Author
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Colas I, Shaw P, Prieto P, Wanous M, Spielmeyer W, Mago R, and Moore G
- Subjects
- Heterochromatin ultrastructure, Recombination, Genetic, Chromatin Assembly and Disassembly, Chromosome Pairing, Chromosomes, Plant genetics, Heterochromatin metabolism, Triticum genetics
- Abstract
During meiosis, homologous chromosomes (homologues) recognize each other and then intimately associate. Studies exploiting species with large chromosomes reveal that chromatin is remodeled at the onset of meiosis before this intimate association. However, little is known about the effect the remodeling has on pairing. We show here in wheat that chromatin remodeling of homologues can only occur if they are identical or nearly identical. Moreover, a failure to undergo remodeling results in reduced pairing between the homologues. Thus, chromatin remodeling at the onset of meiosis enables the chromosomes to become competent to pair and recombine efficiently.
- Published
- 2008
- Full Text
- View/download PDF
30. Molecular characterization of Ph1 as a major chromosome pairing locus in polyploid wheat.
- Author
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Griffiths S, Sharp R, Foote TN, Bertin I, Wanous M, Reader S, Colas I, and Moore G
- Subjects
- Genes, Plant genetics, Heterochromatin genetics, Meiosis genetics, Chromosome Pairing genetics, Chromosomes, Plant genetics, Polyploidy, Triticum genetics
- Abstract
The foundation of western civilization owes much to the high fertility of bread wheat, which results from the stability of its polyploid genome. Despite possessing multiple sets of related chromosomes, hexaploid (bread) and tetraploid (pasta) wheat both behave as diploids at meiosis. Correct pairing of homologous chromosomes is controlled by the Ph1 locus. In wheat hybrids, Ph1 prevents pairing between related chromosomes. Lack of Ph1 activity in diploid relatives of wheat suggests that Ph1 arose on polyploidization. Absence of phenotypic variation, apart from dosage effects, and the failure of ethylmethane sulphonate treatment to yield mutants, indicates that Ph1 has a complex structure. Here we have localized Ph1 to a 2.5-megabase interstitial region of wheat chromosome 5B containing a structure consisting of a segment of subtelomeric heterochromatin that inserted into a cluster of cdc2-related genes after polyploidization. The correlation of the presence of this structure with Ph1 activity in related species, and the involvement of heterochromatin with Ph1 (ref. 6) and cdc2 genes with meiosis, makes the structure a good candidate for the Ph1 locus.
- Published
- 2006
- Full Text
- View/download PDF
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