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The Structural Analysis of Large Noncovalent Oxygen Binding Proteins Current Protein and Peptide Science, 2008, Vol. 9, No. 2 171 Fig. (7). MALLS and ESI-MS analysis of dissociated AmHb. (a,b) Partially dissociated AmHb analyzed by ESI-MS under non-denaturing conditions (a) and by SEC-MALLS (b). (a) The smoothed and subtracted multiply charged spectrum of partially dissociated AmHb, analyzed under non-denaturing conditions. The molecular weight of the components A, B and C were respectively 229 399.22, 70 387.7 and 449 558.7 Da calculated with MaxEnt. The number after the colon in the Inset indicates the number of charges on the ion. The components A were attributed to the 1/12th subunit (D+L), B to a transitory dissociated product and C to truncated HBL-Hb. MaxEnt does not allow the calculation of the other components because of low resolution. They could correspond to other truncated HBL-Hb. The envelop at ~22 000 m/z could correspond to the native HBL-Hb. (b) The SEC-MALLS chromatogram of partially dissociated AmHb allow the observation of native AmHb HBL-Hb at 3608 kDa, of the truncated HBL-Hb and of the 1/12th subunits at 228.8 kDa. (c) The smoothed and subtracted spectrum of AmHb analysed under non-denaturing conditions. Three envelops are observed and MaxEnt allowed the calculation of the component A: 203 660 Da and B: 399 419 Da which correspond respectively to AmHb Dodecamer (D) and dimerisation of D probably due to the high concentration used for the analysis. The complementarities of these data allowed to propose a novel dissociation mechanism for AmHb, through the formation of truncated HBL-Hb, 1/12th subunits and the dissociation of linkers from the dodecamer D [6]. 92
172 Current Protein and Peptide Science, 2008, Vol. 9, No. 2 Bruneaux et al. (a) 4000 Lt y = 1,807x - 2318 R 2 = 0,8485 MALLS mass (kDa) 3800 3600 Rp Md Am Ms Am Ac Ap Pg Pp Lt y = 1,0576x + 46,043 R 2 = 0,9582 3400 Tj y = 1,1252x - 304,64 R 2 = 0,9971 Ev 3200 3200 3400 3600 3800 Model mass (kDa) 4000 (b) 16 Group 3 14 ∆(MALLS mass – model mass) % 12 10 8 6 4 2 (12) Group 1 (6) Group 2 0 Fig. (8). Self-association properties of AmHb. Self-association properties of AmHb followed by MALLS detector during the elution on a gel exclusion column (Superose 6-C). The solid curve represents the refractive index (RI) profile overlaid with the dotted curve which represents the light scattering profil at 90° (LS), versus the retention time. The red curves set for AmHb after dissociation and the blue curves set for AmHb after reassociation. (a) AmHb immediately after exposure in 0.1 M Tris-HCl buffer pH 8.0 and immediately reassociated. (b) AmHb dissociated immediately after exposure in 0.1 M Tris-HCl buffer pH 9.0 and immediately reassociated. (c) AmHb dissociated in 0.1 M Tris-HCl buffer pH 8.0 after 1 hour and reassociated. The percentages of HBL are indicated after dissociation (red) and after reassociation (blue). They are calculated from the integration of HPLC chromatogram at 414 nm. 7. INVESTIGATING RESPIRATORY PIGMENT DI- VERSITY AMONG PHYLA 7.1. HBL-Hb Diversity in Annelids Annelid HBL-Hbs share the same global quaternary structure, as demonstrated by the typical shape observed in electron microscopy for these molecules. However, precise structural studies and three-dimensional reconstructions of their volume in the recent years shed light on high diversity among the different groups. Table 1 presents available data - 2 Ev Cm Tj Ac Am Cm Lt Rp Ms Pp Am Pg Ap Md Lt Species Fig. (9). Comparison between native mass (MALLS) and model mass (ESI-MS). Comparison between measured native mass determined by MALLS and model mass calculated from ESI-MS data. The masses are taken from in Table 1. (a) Plot of MALLS mass versus model mass for annelid HBL-Hbs. The dashed line corresponds to equality between MALLS and model masses. The three linear regression displayed were performed on each group mentioned in (b). (b) Relative difference between MALLS and model masses. The species are sorted by increasing difference value. The values for Carcinus maenas dodecamers (12) and hexamers (6) are added for comparison. Three groups were considered based on the difference values. Ac, Alvinella caudata; Am, Arenicola marina; Ap, Alvinella pompejana; Cm, Carcinus maenas; Ev, Eudistylia vancouverii; Lt, Lumbricus terrestris; Md, Macrobdella decora; Ms, Marphysa sp.; Pg, Paralvinella grasslei; Pp, Paralvinella palmiformis; Rp, Riftia pachyptila. concerning subunit composition of HBL-Hbs from different species. Denaturing ESI-MS is very useful for subunit diversity investigation. It is worthwhile to note that the MS analyses were often performed on a pool from several individuals. Hence the observed diversity is rather representative of the pool and single individuals could exhibit lesser diversity. All 93
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THESE DE DOCTORAT DE L’UNIVERSITE
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i Résumé / Abstract Réponse adap
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iv SOMMAIRE 4.2 Objectifs de l’é
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2 CHAPITRE 1. INTRODUCTION Les éco
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4 CHAPITRE 1. INTRODUCTION 1.1 Mili
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6 CHAPITRE 1. INTRODUCTION Les prin
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8 CHAPITRE 1. INTRODUCTION TAB. 1.1
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10 CHAPITRE 1. INTRODUCTION et cycl
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FIG. 1.5 - Localisation des princip
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(a) FIG. 1.6 - Fonctionnement d’u
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16 CHAPITRE 1. INTRODUCTION TAB. 1.
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18 CHAPITRE 1. INTRODUCTION FIG. 1.
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22 CHAPITRE 1. INTRODUCTION FIG. 1.
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24 CHAPITRE 1. INTRODUCTION le mili
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26 CHAPITRE 1. INTRODUCTION Capacit
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28 CHAPITRE 1. INTRODUCTION TAB. 1.
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30 CHAPITRE 1. INTRODUCTION 700 600
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32 CHAPITRE 1. INTRODUCTION férent
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34 CHAPITRE 1. INTRODUCTION (a) (b)
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40 CHAPITRE 1. INTRODUCTION 1 P 50
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142 CHAPITRE 4. PLASTICITÉ PHÉNOT
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Dénaturant Natif
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172 CHAPITRE 5. ADAPTATIONS RESPIRA
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216 CHAPITRE 6. CONCLUSION ET PERSP
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Annexe A Détermination des masses
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229 10000 h280 .M h) h280−h337 80
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Annexe B Détermination des distrib
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233 Modèle 2 : 2 sous-unités lég
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235 1 0.9 0.8 6 s-u aléatoires 2 s
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Bibliographie Adair, G. S. The hemo
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BIBLIOGRAPHIE 239 Catalina, M. I.,
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BIBLIOGRAPHIE 241 Farmer, C. S., J.
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BIBLIOGRAPHIE 243 Hourdez, S. Adapt
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BIBLIOGRAPHIE 245 Mangum, C. P. Sub
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BIBLIOGRAPHIE 247 Rainer, J., C. P.
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BIBLIOGRAPHIE 249 Svedberg, T. et I
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BIBLIOGRAPHIE 251 Vanin, S., E. Neg
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Liste des figures 1.1 Cycle des mar
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LISTE DES FIGURES 255 4.17 Profils
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Table des matières Résumé Sommai
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TABLE DES MATIÈRES 261 4.3.5 Spect
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TABLE DES MATIÈRES 263 Table des m
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