45 research outputs found

    Divergent selection for fat index in Pannon Ka rabbits: genetic parameters, selection response

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    [EN] The objective of this study was to estimate the response to selection for total body fat content of rabbits measured by computer tomography (CT). A divergent selection experiment was performed using Pannon Ka rabbits, which were previously selected for number of kits born alive. The so-called zero generation consisted of 351 Pannon Ka rabbits, from which the index, total body fat volume (cm3) divided by the body weight (kg), was measured. Rabbits with low and high fat index values were selected to form the parent groups of the lean and fat lines, respectively. The lines consisted of 55-72 females and 35-47 males, depending on the line and generation. After three generations, the rabbits were evaluated by means of a single trait animal model. The fat index showed a moderate heritability estimate (0.28±0.03). The magnitude of the common litter effect was small (0.10±0.02). The breeding values averaged per generation provided slightly asymmetrical responses. Based on the results, the divergent selection was successful in confirming that CT is a very suitable method for performing selection for body composition traits.EFOP-3.6.3-VEKOP-16-2017-00008 project. The project is co-financed by the European Union and the European Social Fund and the JĂĄnos Bolyai Research Scholarship of the Hungarian Academy of Sciences (BO/00871/19)Kasza, R.; Matics, Z.; GerencsĂ©r, Z.; DonkĂł, T.; Radnai, I.; SzendrƑ, Z.; Nagy, I. (2020). Divergent selection for fat index in Pannon Ka rabbits: genetic parameters, selection response. World Rabbit Science. 28(3):129-133. https://doi.org/10.4995/wrs.2020.12733OJS129133283Al-Saef A.M., Khalil M.H., Al-Dobaib S.N., Al-Homidan A.H., GarcĂ­a M.L., Baselga M. 2008. Comparing Saudi synthetic lines of rabbits with the founder breeds for carcass, lean composition and meat quality traits. Livest. Res. Rural Dev., 20: 1-12.DonkĂł T., CzakĂł B., KovĂĄcs Gy., PetnehĂĄzy Ö., Kasza R., SzendrƑ Zs., Garamvölgyi R., Matics Zs. 2016. Total body fat content determination by means of computed tomography (CT) in rabbits. In: Proceedings of the 11th World Rabbit Congress, 16-18 June 2016, Qingdao, China, pp. 753-756.Fortun-Lamothe L. 2006. Energy balance and reproductive performance in rabbit does. Anim. Reprod. Sci., 93: 1-15. https://doi.org/10.1016/j.anireprosci.2005.06.009Garreau H., Eady S.J., Hurtaud J., Legarra A. 2008. Genetic parameters of production traits and resistance to digestive disorders in a commercial rabbit population. In: Xiccato G., Trocino A., Lukefahr S. (eds.) In Proc.: 9th World Rabbit Congress. Fondazione Iniziative Zooprofilattiche e Zootechniche, Verona, Italy, pp. 103-108.Falconer D.S., Mackay T.F.C. 1996. Introduction to Quantitative Genetics. 4th Ed. Longman, London, UK. 1-464.Garreau H., Larzul C., Tudela F., Ruesche J., Ducqrocq V., Fortun-Lamothe L. 2017. Energy balance and body reserves in rabbit females selected for longevity. World Rabbit Sci., 25: 205-213. https://doi.org/10.4995/wrs.2017.5216Groeneveld E. 1990. PEST Users' Manual. Institute of Animal Husbandry and Animal Behaviour Federal Research Centre, Neustadt, Germany 1-61.Groeneveld E., Kovac M., Mielenz N. 2008. VCE User's Guide and Reference manual. Version 6.0. Institute of Farm Animal Genetics, Neustadt, Germany, 1-125.Larzul C., de Rochambeau H. 2005. Selection for residual feed consumption in the rabbit. Livest. Prod. Sci., 95: 67-72. https://doi.org/10.1016/j.livprodsci.2004.12.007Larzul C., Gondret F., Combes S., de Rochambeau H. 2005. Divergent selection on 63-day body weight in the rabbit: response on growth, carcass and muscle traits. Genet. Sel. Evol., 37: 105-122. https://doi.org/10.1051/gse:2004038MartĂ­nez-Álvaro M., HernĂĄndez P., Blasco A. 2016. Divergent selection on intramuscular fat in rabbits: Responses to selection and genetic parameters. J. Anim. Sci., 94: 4993-5003. https://doi.org/10.2527/jas.2016-0590Matics Zs., Nagy I., GerencsĂ©r Zs., Radnai I., Gyovai P., DonkĂł T., Dalle Zotte A., Curik I., SzendrƑ Zs. 2014. Pannon breeding program in rabbit at KaposvĂĄr University. World Rabbit Sci., 22: 287-300. https://doi.org/10.4995/wrs.2014.1511Milisits G., RomvĂĄri R., Dalle Zotte A., SzendrƑ Zs. 1999. Non-invasive study of changes in body composition in rabbits during pregnancy using X-ray computerized tomography. Ann. Zootech., 48: 25-34. https://doi.org/10.1051/animres:19990103Nagy I., Ibåñez N., Mekkawy W., Metzger Sz., Horn P., SzendrƑ Zs. 2006. Genetic parameters of growth and in vivo computerized tomography based carcass traits in Pannon White rabbits. Livest. Sci., 104: 46-52. https://doi.org/10.1016/j.livsci.2006.03.009RomvĂĄri R., Milisits G., SzendrƑ Zs., SĂžrensen P. 1996. Non invasive method to study the body composition of rabbits by X-ray computerized tomography. World Rabbit Sci., 4: 219-224. https://doi.org/10.4995/wrs.1996.298Rouvier R. 1970. VariabilitĂ© gĂ©nĂ©tique du rendement a l'abattage et de la composition anatomique de lapins de trois races. Ann Genet. Sel. Anim., 2: 325-346. https://doi.org/10.1186/1297-9686-2-3-325Shemeis A., Abdallah O.Y. 2000. Possibilities of developing favourable body fat partition via selection indexes - application on rabbits. Arch. Anim. Breed., 43: 193-202. https://doi.org/10.5194/aab-43-193-2000SzendrƑ Zs., RomvĂĄri R., Horn P., Radnai I., BĂ­rĂł-NĂ©meth E., Milisits G. 1996. Two-way selection for carcass traits by computerised tomography. In: Proc. 6th World Rabbit Congress, Toulouse, 2, 371-375.SzendrƑ Zs., Metzger Sz., Nagy I., SzabĂł A., PetrĂĄsi Zs., DonkĂł T., Horn P. 2012. Effect of divergent selection for the computer tomography measured thigh muscle volume on productive and carcass traits of growing rabbits. Livest. Sci., 149: 167-172. https://doi.org/10.1016/j.livsci.2012.07.011Zomeño C., HernĂĄndez P., Blasco A. 2013. Divergent selection for intramuscular fat content in rabbits. 1. Direct response to selection. J. Anim. Sci., 91: 4526-4531. https://doi.org/10.2527/jas.2013-636

    Effect of lighting programme and nursing method on the production and nursing behaviour of rabbit does

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    [EN] The purpose of the experiment was to analyse how the production and nursing behaviour of rabbit does are influenced by different lighting programmes and nursing methods. Rabbit does (n=119) were housed in two rooms. The lighting schedules were a continuous 16L:8D (16L, n=55) or an interrupted 8L:4D:8L:4D (8+8L, n=64). In both rooms, half of the does nursed their kits freely (FS, n=53), while for the other half the suckling method was changed to controlled nursing 3 days prior to the artificial insemination (AI) at day 11 (FS-CS, n=66). Lighting schedule had no significant effect on any productive trait. 76% of the 16L does nursed their kits during the dark period; however, in the 8+8L group, 50% of the nursing events occurred in the dark, 50% during the light periods, respectively. Thus the intermittent lighting disturbed the nursing behaviour of the does. The nursing method significantly affected several traits. AI/parturition, body weight of the does at kindling, number of kits born alive, litter weight at day 21, and suckling mortality were 1.38 and 1.24 (P<0.05), 4.51 and 4.37 kg (P<0.01), 7.95 and 8.46 (P<0.05), 3.06 and 2.92 kg (P<0.05), and 5.3 and 7.3% (P<0.05) in the FS and FS-CS groups, respectively. Compared to the FS group, the advantage of the FS-CS group (P<0.001) was 16.2, 18.4, 9.3 and, 6.3% for total number of kits born, number of kits born alive, number of kits at day 21, and total kitsÂż weight at day 21 per AI, respectively. Due to the change in the nursing method, the frequency of multiple nursing increased. The length of the nursing period of the FS-CS group was significantly exceeded by that of the FS does. Based on these results, changing the nursing method can be used as an adequate biostimulation method.Financial help from TECH_08_A3/2-2008-0384, NDA (National Development Agency) is gratefully acknowledged.GerencsĂ©r, Z.; Matics, Z.; Nagy, I.; Radnai, I.; SzendrĂ”, É.; SzendrĂ”, Z. (2012). Effect of lighting programme and nursing method on the production and nursing behaviour of rabbit does. World Rabbit Science. 20(2):103-116. doi:10.4995/wrs.2012.1078SWORD10311620

    Effect of adult weight and CT-based selection on the performances of growing rabbits

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    The aim of the study was to compare the productive performance of different genotypes. Maternal (M; n=32, adult weight /AW/ 4.0-4.5kg, selected for number of kits born alive), Pannon White (P; n=32, AW: 4.3-4.8kg), and Large body line (L; n=32, AW: 4.8- 5.4kg) (P and L were selected for carcass traits based on CT /Computer tomography/data) rabbits were analysed. Average daily gain between 5-11wk of age, body weight at 11wk of age and feed intake were significantly (P<0.001) highest for L rabbits. For M, P and L rabbits, the following values were observed: average daily gain=38.6, 43.1 and 47.4g/d; body weight=2458, 2667 and 2949g; feed intake=115, 121 and 138g/d, respectively. Mortality of growing rabbits was unaffected by genotype. It can be concluded that production traits were mainly affected by the adult weight of the genotypes

    Effect of hair shearing on live performance and carcass traits of growing rabbits under hot ambient temperature

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    [EN] The aim of the study was to examine the effect of hair shearing in growing rabbits reared at high ambient temperature. The live performance and carcass traits of growing rabbits reared at 20°C (not sheared, C, n=50) or at 28°C (not sheared, H, n=50, or sheared at 5, 7 and 9 wk, HS, n=50) were compared. The ambient temperature and relative humidity were 20.5±1.1°C and 54±11% in the 20°C room and 28.8±0.2°C and 35±8% in 28°C room, respectively. Feed intake of H and HS groups decreased by 29.0 and 20.4%, respectively, compared to C rabbits (P&lt;0.001). The same data for weight gain were 24.6 and 16.9% (P&lt;0.001), and for body weight at 12 wk were 16.8 and 11.5% (P&lt;0.001). At the same time, the feed conversion ratio improved (C: 3.53, HS: 3.34, H: 3.31; P&lt;0.001). Nevertheless, the mortality rate of rabbits was not affected by the studied treatment and was overall low (0-4%). No differences were observed in dressing out percentages either (ratio of chilled carcass (CC) to the slaughter weight: 61.6-61.9%). The ratio of liver to CC differed among the experimental groups, with the highest value recorded in C group and the lowest in H group; HS rabbits showed intermediate results (C: 4.86%, HS: 4.27%, H: 3.91%; P&lt;0.001). Lower ratios of fat deposits to reference carcass were also observed in rabbits kept at high ambient temperature (perirenal fat: C: 2.59%, HS: 1.82%, H: 1.60%; P&lt;0.001; scapular fat: C: 0.89%, HS: 0.66%, H: 0.51%; P&lt;0.001). It can be concluded that the negative effect of higher ambient temperature (28 vs. 20°C) on production in growing rabbits can be reduced significantly by hair shearing.En este agradecimieento: "The work was supported by the GINOP-2.3.4-15-2016-00005 project. Publication was supported by the EFOP-3.6.3-VEKOP-16–2017–00008 project. The project is co-funded by the European Union and the European Social Fund"Matics, Z.; Kasza, R.; GerencsĂ©r, Z.; Radnai, I.; Dalle Zotte, A.; Cullere, M.; SzendrƑ, Z. (2020). Effect of hair shearing on live performance and carcass traits of growing rabbits under hot ambient temperature. World Rabbit Science. 28(3):161-167. https://doi.org/10.4995/wrs.2020.13164OJS161167283Balnave D. 1972. The effect of temperature and length of exposure on liver composition and hepatic lipogenic enzyme activity in the immature male chick (Gallus domesticus). Comp. Biochem. Physiol., 438: 999-1007. https://doi.org/10.1016/0305-0491(72)90244-1Blasco A., Ouhayoun J. 1996. Harmonization of criteria and terminology in rabbit meat research. Revised proposal. World Rabbit Sci., 4: 93-99. https://doi.org/10.4995/wrs.1996.278Chiericato G.M., Rizzi C., Rostellato V. 1993. Effect of genotype and environmental temperature on performance of the young meat rabbit. World Rabbit Sci., 1: 119-125. https://doi.org/10.4995/wrs.1993.204Chiericato G.M., Ravarotto L., Rizzi R. 1994. Study of the metabolic profile of rabbits in relation to two different environmental temperatures. World Rabbit Sci., 2: 153-160. https://doi.org/10.4995/wrs.1994.232Chiericato G.M., Rizzi C., Rostellato V. 1996. Growth and slaughtering performance of three rabbit genotypes under different environmental conditions. Ann. Zootech., 45: 311-318. https://doi.org/10.1051/animres:19960403Deltoro J., LĂłpez A.M. 1986. Development of commercial characteristics of rabbit carcasses during growth. Livest. Prod. Sci., 15: 271-283. https://doi.org/10.1016/0301-6226(86)90034-5EC 2010. Directive 2010/63/EU of the European Parliament and of the Council of 22 September 2010 on the protection of animals used for scientific purposes. Official Journal of the European Union L276: 33-79.FernĂĄndez-Carmona J., Cervera C., Sabater C., Blas E. 1995. Effect of diet composition on the production of rabbit breeding does housed in a traditional building and at 30°C. Anim. Feed Sci. Technol., 52: 289-297. https://doi.org/10.1016/0377-8401(94)00715-LFinzi A., Morera P., Kuzminsky G. 1992. Effect of shearing on rabbit bucks performances in hot ambient conditions. J. Appl. Rabbit Res., 15: 489-494.Fuquay J.W. 1981. Heat stress as it affects animal production. J. Anim. Sci., 52: 164-174. https://doi.org/10.2527/jas1981.521164xHermes I.H., Ahmed B.M., Khalil M.H., Salah M.S., Al-Homidan A.A. 1999. Growth performance, nutrients utilization and carcass traits of growing Californian rabbits raised under different ambient temperatures. Egypt. J. Rabbit Sci., 9: 117-138.Jackson R., Rogers A.D, Lukefahr S.D. 2006. Effects of the naked gene on postweaning performance and thermotolerance characters in fryer rabbits: Final results. World Rabbit Sci., 14: 147-155. https://doi.org/10.4995/wrs.2006.559Kovitvadhi A., Chundang P., Thongprajukaew K., Tirawattanawanich C. 2019. Effects of different ambient temperatures on growth performances, digestibility, carcass traits and meat chemical components in fattening rabbits. J. Agriculture, 35: 495-502.Lebas F., Ouhayoun J. 1987. Incidence du niveau protĂ©ique de l'aliment, de milieu d'Ă©levage et de la saison sur la croissance et les qualitĂ©s bouchĂ©res du lapin. Ann. Zootech., 36: 421-432. https://doi.org/10.1051/animres:19870406Lebas F., Coudert P., de Rochambeau H., ThĂ©bault R.G. 1997. The rabbit: husbandry, health and production. FAO Anim. Prod. and Health Series No. 21Lukefahr S.D., Ruiz-Feria C.A. 2003. Rabbit growth performance in a subtropical and semi-arid environment: Effects of fur clipping, ear length, and body temperature. Livest. Res. Rural Devel. 15: 2. Available at http://www.cipav.org.co/lrrd/lrrd15/2/luke152.htm Accessed October 2019.Marai I.F.M., Habeeb A.A.M., Gad A.E. 2002. Rabbits' productive, reproductive and physiological performance traits as affected by heat stress: a review. Livest. Prod. Sci., 78: 71-90. https://doi.org/10.1016/S0301-6226(02)00091-XMaya-Soriano M.J., Taberner E., Sabes-Alsina M., Ramon J., Rafel O., Tusell L., Piles M., LĂłpez-BĂ©jar M. 2015. Daily exposure to summer temperatures affects the motile subpopulation structure of epididymal sperm cells but not male fertility in an in vivo rabbit model. Theriogenology, 84: 384-389. https://doi.org/10.1016/j.theriogenology.2015.03.033Metzger Sz. 2006. Examination on carcass traits and meat quality of rabbit. (in Hung.) Doctoral (Ph.D.) dissertation. pp. 135.NASA https://climate.nasa.gov/Perez J.M., Lebas F., Gidenne T., Maertens L., Xiccato G., Parigi-Bini R., Dalle Zotte A., Cossu M.E., Carazzolo A., Villamide M.J., Carabaño R., Fraga M.J., Ramos M.A., Cervera C., Blas E., FernĂĄndez J., FalcĂŁo-e-Cunha L., Bengala Freire J. 1995. European reference method for in vivo determination of diet digestibility in rabbits. World Rabbit Sci. 3: 41-43. https://doi.org/10.4995/wrs.1995.239Renaudeau D., Collin A., Yahav S., de Basilio V., Gourdine J.L., Collier R.J. 2012. Adaptation to hot climate and strategies to alleviate heat stress in livestock production. Animal, 6: 707-728. https://doi.org/10.1017/S1751731111002448SAS Version 9.4. 2014. SAS Institute Inc; Cary, NC. Schlolaut W. 1995. Das grosse Buch vom Kaninchen. DLG-Verlag, Frankfurt am Main.Stephan E. 1980. The influence of environmental temperatures on meat rabbits of different breeds. Commercial Rabbit, 8: 12-15.SzendrƑ Zs., Rashwan R.R., BirĂł-NĂ©meth E., Radnai I., Orova Z. 2007. Effect of shearing of hair in summer on production of rabbit does. Acta Agr. Kapos., 11: 37-42.SzendrƑ Zs., Papp Z., Kustos K. 2018. Effect of ambient temperature and restricted feeding on the production of rabbit does and their kits. Acta Agr. Kapos., 22: 1-17. https://doi.org/10.31914/aak.2272Verga M., Luzi F., Carenzi C., 2007. Effects of husbandry and management systems on physiology and behaviour of farmed and laboratory rabbits. Horm. Behav., 52, 122-129. https://doi.org/10.1016/j.yhbeh.2007.03.024Zeferino P.C., Moura T.M.A.S.A., Fernandes S., Kanayama S.J., Scapinello C., Sartori R.J. 2011. Genetic group × ambient temperature interaction effects on physiological responses and growth performance of rabbits. Livest. Sci., 140: 177-183. https://doi.org/10.1016/j.livsci.2011.03.02

    Food additives: Sodium benzoate, potassium sorbate, azorubine, and tartrazine modify the expression of NFÎșB, GADD45α, and MAPK8 genes

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    It has been reported that some of the food additives may cause sensitization, inflammation of tissues, and potentially risk factors in the development of several chronic diseases. Thus, we hypothesized that expressions of common inflammatory molecules – known to be involved in the development of various inflammatory conditions and cancers – are affected by these food additives. We investigated the effects of commonly used food preservatives and artificial food colorants based on the expressions of NFÎșB, GADD45α, and MAPK8 (JNK1) from the tissues of liver. RNA was isolated based on Trizol protocol and the activation levels were compared between the treated and the control groups. Tartrazine alone could elicit effects on the expressions of NFÎșB (p = 0.013) and MAPK8 (p = 0.022). Azorubine also resulted in apoptosis according to MAPK8 expression (p = 0.009). Preservatives were anti-apoptotic in high dose. Sodium benzoate (from low to high doses) dose-dependently silenced MAPK8 expression (p = 0.004 to p = 0.002). Addition of the two preservatives together elicited significantly greater expression of MAPK8 at half-fold dose (p = 0.002) and at fivefold dose (p = 0.008). This study suggests that some of the food preservatives and colorants can contribute to the activation of inflammatory pathways

    Comparison of productive and carcass traits and economic value of lines selected for different criteria, slaughtered at similar weights

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    [EN] The aim of the experiment was to compare 3 genetic groups, slaughtered at similar weights, to examine their productive and carcass traits and economic value. Three lines of the Pannon Breeding Programme, selected for different criteria, were examined in the experiment. Pannon Ka (PKa, maternal line) does were inseminated with semen of PKa, Pannon White (PWhite) or Pannon Large (PLarge, terminal line) bucks. The kits (PKa×PKa, PWhite×PKa, PLarge×PKa; n=60 in each genetic group) were weaned at 35 d of age and reared until 88, 83 and 79, respectively, when they reached similar body weights for slaughtering (2.8 kg). The weight gain of PLarge×PKa was the largest (51.0 g/d) and that of PKa×PKa was the smallest (47.2 g/d), while PWhite×PKa (41.8 g/d) was intermediate (P<0.001). Difference was found in feed conversion ratio between weaning and the age of slaughter  PKa×PKa: 3.03 respect to PWhite×PKa: 2.75 and PLarge×PKa: 2.66; , P<0.05). Dressing out percentage and ratio of hind part to reference carcass of PWhite×PKa, PLarge×PKa and PKa×PKa were 62.4 and 37.7, 61.8 and 37.5, 61.3 and 36.8%, respectively (P<0.01). Results show that PLarge×PKa rabbits were able to exceed the average economic indicators compared to other groups. It may be concluded that the production performance of growing rabbits was affected by the adult weight, but the carcass traits were influenced by the computer tomography (CT)-based selection.This paper was supported by the JĂĄnos Bolyai Research Scholarship of the Hungarian Academy of Sciences, and by the GOP-1.1.1-11-2012-0132 project.SzendrƑ, K.; SzendrƑ, Z.; GerencsĂ©r, Z.; Radnai, I.; Horn, P.; Matics, Z. (2016). Comparison of productive and carcass traits and economic value of lines selected for different criteria, slaughtered at similar weights. World Rabbit Science. 24(1):15-23. https://doi.org/10.4995/wrs.2016.3684152324

    The ALICE detector data link

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    The ALICE detector data link has been designed to cover all the needs for data transfer between the detector and the data-acquisition system. It is a 1 Gbit/s, full-duplex, multi-purpose fibre optic link that can be used as a medium for the bi-directional transmission of data blocks between the front-end electronics and the data- acquisition system and also for the remote control and test of the front-end electronics, In this paper the concept, the protocol, the specific test tools, the prototypes of the detector data link and the read-out receiver card, their application in the ALICE-TPC test system and the integration with the DATE software are presented. The test results on the performance are also shown. (14 refs)

    Finite Difference Schemes for Stochastic Partial Differential Equations in Sobolev Spaces

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    We discuss LpL_p-estimates for finite difference schemes approximating parabolic, possibly degenerate, SPDEs, with initial conditions from WpmW^m_p and free terms taking values in Wpm.W^m_p. Consequences of these estimates include an asymptotic expansion of the error, allowing the acceleration of the approximation by Richardson's method.Comment: 22 pages. The final publication is available at Springer via http://dx.doi.org/10.1007/s00245-014-9272-

    Electron Transfer from Cyt b559 and Tyrosine-D to the S2 and S3 states of the water oxidizing complex in Photosystem II at Cryogenic Temperatures

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    The Mn4CaO5 cluster of photosystem II (PSII) catalyzes the oxidation of water to molecular oxygen through the light-driven redox S-cycle. The water oxidizing complex (WOC) forms a triad with Tyrosine(Z) and P-680, which mediates electrons from water towards the acceptor side of PSII. Under certain conditions two other redox-active components, Tyrosine(D) (Y-D) and Cytochrome b (559) (Cyt b (559)) can also interact with the S-states. In the present work we investigate the electron transfer from Cyt b (559) and Y-D to the S-2 and S-3 states at 195 K. First, Y-D (aEuro cent) and Cyt b (559) were chemically reduced. The S-2 and S-3 states were then achieved by application of one or two laser flashes, respectively, on samples stabilized in the S-1 state. EPR signals of the WOC (the S-2-state multiline signal, ML-S-2), Y-D (aEuro cent) and oxidized Cyt b (559) were simultaneously detected during a prolonged dark incubation at 195 K. During 163 days of incubation a large fraction of the S-2 population decayed to S-1 in the S-2 samples by following a single exponential decay. Differently, S-3 samples showed an initial increase in the ML-S-2 intensity (due to S-3 to S-2 conversion) and a subsequent slow decay due to S-2 to S-1 conversion. In both cases, only a minor oxidation of Y-D was observed. In contrast, the signal intensity of the oxidized Cyt b (559) showed a two-fold increase in both the S-2 and S-3 samples. The electron donation from Cyt b (559) was much more efficient to the S-2 state than to the S-3 state
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