ZHU Yanbing, LI Hebin, NI Hui, LIU Jingwen, XIAO Anfeng, CAI Huinong. Purification and biochemical characterization of manganesecontaining superoxide dismutase from deep-sea thermophile Geobacillus sp. EPT3[J]. Acta Oceanologica Sinica, 2014, 33(12): 163-169. doi: 10.1007/s13131-014-0534-2
Citation: ZHU Yanbing, LI Hebin, NI Hui, LIU Jingwen, XIAO Anfeng, CAI Huinong. Purification and biochemical characterization of manganesecontaining superoxide dismutase from deep-sea thermophile Geobacillus sp. EPT3[J]. Acta Oceanologica Sinica, 2014, 33(12): 163-169. doi: 10.1007/s13131-014-0534-2

Purification and biochemical characterization of manganesecontaining superoxide dismutase from deep-sea thermophile Geobacillus sp. EPT3

doi: 10.1007/s13131-014-0534-2
  • Received Date: 2013-03-15
  • Rev Recd Date: 2013-09-13
  • Thermostable SOD is a promising enzyme in biotechnological applications. In the present study, thermophile Geobacillus sp. EPT3 was isolated from a deep-sea hydrothermal field in the East Pacific. A thermostable superoxide dismutase (SOD) from this strain was purified to homogeneity by steps of fractional ammonium sulfate precipitation, DEAE-Sepharose chromatography, and Phenyl-Sepharose chromatography. SOD was purified 13.4 fold to homogeneity with a specific activity of 3 354 U/mg and 11.1% recovery. SOD from Geobacillus sp. EPT3 was of the Mn-SOD type, judged by the insensitivity of the enzyme to both KCN and H2O2. SOD was determined to be a homodimer with monomeric molecular mass of 26.0 kDa. It had high thermostability at 50℃ and 60℃. At tested conditions, SOD was relatively stable in the presence of some inhibitors and denaturants, such as β-mercaptoethanol (β-ME), dithiothreitol (DTT), phenylmethylsulfonyl fluoride (PMSF), urea, and guanidine hydrochloride. Geobacillus sp. EPT3 SOD showed striking stability across a wide pH range from 5.0 to 11.0. It could withstand denaturants of extremely acidic and alkaline conditions, which makes it useful in the industrial applications.
  • Magmatism at mid-ocean ridges is one of our planet’s most important geological processes as it forms the oceanic crust, which covers nearly two-thirds of the Earth’s surface. Unlike fast spreading ridges, it is widely accepted that ultraslow–spreading ridges have relatively low extents of melting and magmatism, thinner crust, and experience limited crustal processes (Dick, 1989). Although ultraslow–spreading ridges are characterized by low magma supplies, there is also evidence of substantial magmatic processes in magma reservoirs, rather than the stable magma chambers present under some segments (Dick et al., 2003; Coogan et al., 2001; Jian et al., 2017). Until now, how magmas behave in the crust and how the thermal and dynamic regimes of the magma reservoirs and conduit systems effect the eruption mechanism of magmas from ultraslow mid-ocean ridges (MORs) has received comparatively little attention. As the product of erupted magma, mid-ocean ridge basalts (MORBs) can provide information about the magmatism under ridges (Yang et al., 2013). Therefore, unraveling the chemical effects of magmatic processes reflected by the MORBs is key to determining the details of the magma plumbing system beneath ultraslow oceanic spreading centers.

    Previous studies have mainly focused on the whole–rock geochemical and isotopic compositions of MORBs from ultraslow–spreading ridges (Dick et al., 2003; Yang et al., 2013, 2017). However, whole–rock geochemistry cannot provide information about the magmatic processes occurring within the crust, which we are attempting to understand, because the bulk–rock compositions of MORBs reflect the variable overprinting of the low–pressure differentiation of mantle–derived primary liquids (Stolper, 1980; Yang et al., 2013). Unlike whole–rock compositions, the major and trace element contents of plagioclase phenocrysts are easily preserved and are sensitive to the physicochemical conditions of the melt from which they crystallized (Ginibre et al., 2004; Mollo et al., 2011; Mutch et al., 2019; Bennett et al., 2019). Their geochemical compositions and morphology are commonly used to obtain information about the conditions of the magmatic system from which they crystallized. For example, Mutch et al. (2019) established an element diffusion model for plagioclase to constrain the timescales of magmatic processes. Bennett et al. (2019) demonstrated that various plagioclase textures in mid–ocean ridge basalts can be used as indicators of various magmatic processes. In addition, the composition of plagioclase can provide information about the conditions of the magmatic system, such as the water content, temperature, and melt composition (Hellevang and Pedersen, 2008; Lange et al., 2013; Coote and Shane, 2016). These conditions are related to magma mixing, fractional crystallization, and assimilation processes. Therefore, plagioclase phenocrysts have the potential to record a magmatic history that might be obscured at the whole–rock geochemical scale. However, compared with continental and other oceanic settings, plagioclase phenocrysts are still underutilized in the study of MORBs from ultraslow spreading ridges.

    In recent years, the China Ocean Mineral Resources R&D Association (COMRA) has provided support for scientific expeditions to the Southwest Indian Ridge (SWIR), during which a large number of new samples were collected (Tao et al., 2012). Among the various types of MORBs, plagioclase ultraphyric basalts (PUBs), defined by Cullen et al. (1989), are valuable due to their high plagioclase contents (10%–54%). These plagioclase phenocrysts contain unique information about crustal processes and the characteristics of the magmatic array present in the lower oceanic crust (Hellevang and Pedersen, 2008; Lange et al., 2013; Bennett et al., 2019). In this study, we examined the compositions of plagioclase phenocrysts from PUBs erupted on the SWIR (51°E), which is a typical ultraslow spreading ridge. Microanalysis of these phenocrysts provides an opportunity to investigate the geochemical changes that occurred, and thus, provides insights into the details of the magma reservoirs beneath the ultraslow oceanic spreading centers.

    The SWIR is a typical ultraslow spreading ridge, with a half–spreading rate of around 7–9 mm/a (Dick et al., 2003). It separates the African Plate from the Antarctic Plate. The SWIR extends for 8 000 km from the Rodrigues Triple Junction (RTJ, 70°E) at its eastern end to the Bouvet Triple Junction at its western end (BTJ, 0°) (Fig. 1a). The SWIR is characterized by strong segmentation and discontinuous magmatism. Bathymetric data has revealed a shallow central region between the Prince Edward Transform Fracture Zone (35.5°E) and the Gallieni Fracture Zone (52.3°E), with an average depth of ~3 200 m compared with the deeper western (~4 000 m) and the eastern (~4 500 m) sections of the ridge (Cannat et al., 2008). As a result of the Marion hotspot to the southwest, this region has strong negative residual mantle Bouguer gravity anomalies, indicating relatively active crust–mantle exchange, deep magmatism, moderate levels of melting, and a moderate heat supply (Georgen et al., 2001; Sauter et al., 2009).

    Figure  1.  Bathymetric map of the Southwest Indian Ridge (SWIR) (a) and location of the sample (b) (http://www.geomapapp.org). The location of the sample used in this study is marked by the star. BTJ: Bouvet Triple Junction; RTJ: Rodrigues Triple Junction; PE: Prince Edward Transform Fracture Zone; IFZ: Indomed Fracture Zone; GFZ: Gallieni Fracture Zone.

    The study area and the sample sites are located between the Indomed (46°E) and Gallieni fracture zones (IFZ–GFZ) on the shallow central region of the SWIR (Fig. 1b). Previous geophysical and geochemical studies have been conducted on the ridge segment between the IFZ and GFZ. The center of this segment has anomalously thick crust (up to 10 km) (Niu et al., 2015) compared with that of the neighboring ridge sections. This thick oceanic crust indicates a robust magma supply in this area, which has been inferred to be associate with the Crozet hotspot (Sauter et al., 2009; Zhang et al., 2013) or with tectonic processes (Jian et al., 2017). Segments with robust magma supplies are also promising areas for hydrothermal activity (Tao et al., 2012).

    The PUB sample examined in this study was collected by television–guided grabs (TVGs) during the R/V Dayang Yihao Cruise DY115–21 to the 51°E magmatic segment in 2010. The sample was collected at a water depth of about 1 655 m. Optical microscopy analysis indicates that lavas from the SWIR are porphyritic and contain ~15% phenocrysts. The phenocrysts are mostly euhedral to subhedral plagioclase with polysynthetic twinning. The other main phase is subhedral to anhedral olivine (<1%). The plagioclase crystals contain abundant melt inclusions. The groundmass is primarily composed of suhedral, lath–like plagioclase and subordinate, anhedral olivine (Fig. 2).

    Figure  2.  Photographs and representative photomicrographs (cross–polarized) of the PUB from the SWIR. a. Hand specimens of the sample; b. enhedral to subhedral plagioclase phenocrysts with melt inclusions; and c suhedral. lath–like plagioclase and subordinate, anhedral olive. Pl: plagioclase, Ol: olivine, MI: melt–melt inclusion.

    Backscattered electron (BSE) imaging of the plagioclase in polished thin sections was used to characterize the textures of the crystals. Mineral analysis of the plagioclase was conducted using the JEOL JXA 8100 electron microprobe at the Key Laboratory of Submarine Geosciences (KLSG), Ministry of Natural Resources (MNR). The analytical conditions were as follows: a 15 kV accelerating voltage, a 20 nA specimen current, and a 1 μm focused beam. The peak counting times were 90 s for Fe and Mg, and 20 s for all of the other major elements. The detection limits for most of the elements, except for Ti, were lower than 400×10–6, depending on the abundance of the elements. The detection limit of Ti was 600×10–6 due to its lower content. The following natural and synthetic standards were used for the specified elements: Olivine (Si, Mg), Apatite (Ca, P), Hematite (Fe), Albite (Na, Al), Orthoclase (K), Rhodonite (Mn), Rutile (Ti), and Tugtupite (Cl). The raw data was corrected using the ZAF correction. The chemical formulas of the plagioclase phenocrysts were calculated from the mineral analysis results based on 24 anions.

    The plagioclase phenocrysts (typically >0.5 mm) from the SWIR are predominantly euhedral to subhedral in shape with tabular habits. They exhibit three textural types (Figs 3ac). Type 1 plagioclase crystals are characterized by oscillatory zoning, surrounded by a thin rim (<50 μm). Type 2 plagioclase crystals are also characterized by oscillatory zoning, but have wider rims (50–150 μm) than Type 1 plagioclase (<50 μm). Type 3 plagioclase crystals contain numerous circular melt inclusions and do not exhibit oscillatory zoning. The plagioclase in the groundmass varies in size. The relatively large microphenocrysts typically have sizes of 0.01–0.50 mm and are primarily unzoned, whereas the relatively small groundmass microlites are <0.01 mm (Fig. 3d).

    Figure  3.  Back-scattered-electron (BSE) images of the different types of plagioclase in the PUB. a. Type 1 plagioclase phenocryst; b. Type 2 plagioclase phenocryst; c. Type 3 plagioclase phenocryst; and d. plagioclase microphenocrysts. MI: melt inclusion.

    A total of 25 microprobe analyses were performed on the studied sample. Representative chemical data for the plagioclases are presented in Table 1. These phenocrysts have An contents of 58 to 82. No distinct compositional differences exist among the three types of plagioclase phenocrysts. All of the phenocrysts have calcic cores (An74–82) and sodic rim growth (~An67–71). The variation from the core to the rim is 10–20 mol% An. The plagioclase crystals found within the microphenocrysts and microlites are sodic (An58–63), similar to the rims of the phenocrysts. On the ternary classification diagram, the plagioclase in the studied sample display a continuous range from bytownite to labradorite, with An decreasing from 82 to 58. The cores of the plagioclase phenocrysts are bytownite, while the rims range from bytownite to labradorite. All of the plagioclase crystals in the groundmass are labradorite with lower An contents (Fig. 4).

    Table  1.  Representative microprobe data for the plagioclase from the SWIR of this study
    ElementPhenocrystalGroundmass1)
    Type IType IIType III Pl-mPl-g
    CoreRimCoreRimCoreRim
    SiO248.2551.4250.7352.2448.8051.7453.7054.69
    TiO20.060.100.050.000.000.110.080.17
    Al2O331.4930.0330.2829.6131.7029.8128.4126.72
    FeO0.290.400.500.540.380.620.581.41
    MnO0.040.000.040.000.000.000.000.00
    MgO0.150.190.240.180.200.200.180.37
    CaO16.8014.5214.9414.0116.2914.1312.7711.92
    Na2O1.993.252.923.422.213.224.154.64
    K2O0.010.020.010.030.010.020.040.06
    P2O50.010.030.000.020.010.000.010.05
    Total99.0899.9599.71100.0499.5999.8699.92100.02
    Calculated atoms based on 24 oxygens
    Si6.7047.0316.9627.1276.7367.0727.3117.457
    Ti0.0060.0100.0050.0000.0000.0110.0080.017
    Al5.1574.8404.8994.7615.1574.8034.5594.294
    Fe0.0340.0460.0570.0620.0430.0710.0660.161
    Mn0.0050.0000.0040.0000.0000.0000.0000.000
    Mg0.0310.0390.0490.0360.0410.0410.0370.074
    Ca2.5012.1272.1982.0472.4092.0701.8631.742
    Na0.5360.8620.7760.9050.5910.8521.0961.225
    K0.0020.0030.0020.0050.0010.0030.0070.011
    P0.0020.0030.0000.0020.0010.0000.0010.005
    Total14.97714.96114.95314.94514.98014.92414.94714.987
    An0.820.710.740.690.800.710.630.58
    Ab0.180.290.260.310.200.290.370.41
    Or0.000.000.000.000.000.000.000.01
    FeO/MgO1.912.122.083.041.903.063.163.87
    Ca/Na4.672.472.832.264.072.431.701.42
    T/°C2)12961108123710981280112910631042
    Note: 1) Pl-m and Pl-g represent plagioclase microphenocrysts and microlites in groundmass, respectively; 2) crystallization temperature is calculated according to Kudo and Weill (1983).
     | Show Table
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    Figure  4.  Or–Ab–An classification diagram of plagioclase.

    In terms of the major elements, the FeO and MgO concentrations of the plagioclase do not vary significantly. The FeO and MgO concentrations are 0.29%–1.41% (wt) and 0.15%–0.37% (wt), respectively. The FeO and MgO concentrations of the plagioclase cores are relatively depleted compared with those of the plagioclase rims and groundmass.

    Plagioclase compositions are a useful indicator of crystallization temperature (Kudo and Weill, 1970; Mollo et al., 2011). In this study, the plagioclase–melt geothermometry method proposed by Kudo and Weill (1970) was used to estimate the crystallization temperature of the plagioclase. Before applying the geothermometry method, the pressures must be determined. According to Chen et al. (2002), the pressures of the plagioclase rims and cores are approximately 0.5×108 and 1.0×108 Pa, respectively. Similarly, the plagioclase in the groundmass is estimated to have crystallized at shallower depths, within the upper crust or on the seabed. The pressure of the groundmass is also assumed to be 0.5×108 Pa according to Chen et al. (2002). Based on the assumptions stated above, Eqs (1) and (2) were used to calculate the crystallization temperatures of the plagioclase crystals:

    $$ \frac{{{\rm{ln}}\lambda }}{{{\sigma }}} + 1.29 \times \frac{{{{10}^4}\varphi }}{T} = 9.87 \times {10^{ - 3}}T - 15.21,\left( {{P_{{{\rm{H}}_{\rm{2}}}{\rm{O}}}} = 0.5\times 10^8\;{\rm {Pa}}} \right), $$ (1)
    $$ \frac{{{\rm{ln}}\lambda }}{\sigma } + 1.29 \times \frac{{{{10}^4}\varphi }}{T} = 9.60 \times {10^{ - 3}}T - 15.76,\left( {{P_{{{\rm{H}}_{\rm{2}}}{\rm{O}}}} = 1.0\times 10^8\;{\rm {Pa}} } \right), $$ (2)

    where $ \lambda$=(XNaXSi/XCaXAl) is for the groundmass, $ \sigma$=(XAbγAb/XAnγAn) is for the plagioclase, and $ \varphi$=(XCa+XAlXSiXNa) is for the groundmass. X represents the mole fraction of the component. Although the groundmass is microcrystalline and lacks glass, Lange et al. (2013) demonstrated that PUB hosted glasses have the same range of compositions as aphyric lavas from the same segment. Therefore, in the calculation process, we use the average whole–rock compositions of the aphyric basalts from the same segment, which were reported by Yang et al. (2014), as an approximate proxy for the groundmass.

    According to the plagioclase geothermometer described above, the crystallization temperatures of the phenocryst cores and rims are (1 273±18)°C and (1 099±10)°C, respectively. From the core to the rim of the phenocryst, the crystallization temperature decreases by about 200°C. The crystallization temperatures of the microphenocrysts and microlites in the groundmass range from 1 063°C to 1 087°C (average = 1 072°C), which is similar to the crystallization temperature of the phenocryst rims (Table 1).

    In previous studies, the crystallization temperatures of high–An (An≥70%) plagioclase phenocrysts, which were estimated from the entrapment temperature of melt inclusions in samples from ultraslow spreading ridges, were found to range from 1 230°C to 1 260°C (Nielsen et al., 1995; Drignon et al., 2019). Whereas the minimum estimation of the crystallization temperature of low An (An<70%) plagioclase is 1 100°C (Yang et al., 2019). The crystallization temperatures in our study are consistent with these results, which suggests that our calculated results are reasonable and the plagioclases thermometer by Kudo and Weill (1970) can be used to calculate the crystallization temperatures of plagioclase phenocrysts from MORBs beneath ultraslow ridges.

    In plagioclase, the diffusion rates of major elements, such as the NaSi–CaAl exchange, are extremely slow (Grove et al., 1984). The geochemical zoning of the crystals likely reflects the magmatic conditions, and thus, it can be used to investigate the likelihood of plagioclase–melt equilibrium. Previous experiments have demonstrated that the partition coefficient of Ca/Na (KCa/Na) between plagioclase and melt mainly positively depends on the water content of the magma (Sisson and Grove, 1993; Martel et al., 2006). For basaltic magmas from mid-ocean ridges, when the magmatic water content is 3%, KCa/Na is typically ~1 for mid to upper lithospheric pressures (<10×108 Pa) (Sisson and Grove, 1993). The magmatic water contents of the SWIR lavas are close to the global average value for the upper mantle (0.3%–0.4%) (Robinson et al., 2001), which indicates that their KCa/Na value is less than 1.

    In addition to the KCa/Na value, a representative melt composition is required to assess the equilibrium composition of the plagioclase phenocrysts. The groundmass of these rocks is microcrystalline and lacks glass, so it represents the final melt. Thus, as was discussed above, the average whole–rock composition of the aphyric basalts from the same segment is representative of the groundmass composition. The reported composition of the basalts and glass from this ridge segment do not vary significantly, with molar Ca/Na values ranging from 2.11 to 2.68 (Yang et al., 2014; Bézos and Humler, 2005). The molar Ca/Na values of the plagioclase phenocrysts in equilibrium with the melt are consistently≤2.68 (Fig. 5). The molar Ca/Na values of the plagioclase rims and groundmass range from 1.42 to 2.47, which is in equilibrium with the melt (Fig. 5). However, the compositions of the plagioclase cores (Ca/Na = 3 to 4) are higher than the upper limit of the equilibrium melt (Ca/Na = 2.68) (Fig. 5). In addition, the crystallization temperatures of the plagioclase cores (average of 1 273°C) are close to the experimental melting point of basaltic magma (~1 300°C) and the estimated upper mantle potential temperature of the SWIR lavas (~1 280°C) (Kamenetsky et al., 2000; Robinson et al., 2001). Therefore, the plagioclase cores are unlikely to have formed in the host magma. Instead, they are most likely xenocrysts, which crystallized from a more calcic melt.

    Figure  5.  Ca/Na molecular ratio of the plagioclase compared with that of the melt. The melt represents the range of glass and whole-rock compositions, the data are obtained from Yang et al. (2014) and Bézos and Humler (2005). The K values and water contents are from Martel et al. (2006). Equilibrium between the plagioclase and melts is possible in the shaded area.

    It is worth noting that compared with the plagioclase rims, the plagioclase phenocrysts in the groundmass have lower Ca/Na ratios (<2). Their more sodic compositions are most likely due to decreasing magmatic water contents caused by the fact that the KCa/Na between the plagioclase and the melt decreases as of the water content of the magma increases (Martel and Schmidt, 2003). Water is lost during the late-stage of magma ascent through the conduit due to ascent–driven decompression of the water-saturated magma. Thus, the compositional variations can be explained by the variations in the magmatic water content.

    The MgO zoning patterns of the plagioclase phenocrysts have the potential to record the magmatic composition because the Mg contents of the plagioclase phenocrysts reflect the composition of the host melt (Ginibre et al., 2002). Although it is difficult to accurately estimate the Mg partition coefficient (KMg) between plagioclase and melt, empirical studies have suggested an Arrhenius-like relationship of decreasing KMg with increasing XAn (Bindeman et al., 1998). To investigate the equilibrium relationship between the SWIR plagioclase and the host magma, we estimated the KMg based on this empirical relationship, which is often used to determine whether plagioclase phenocrysts are in equilibrium with the melt in magmatic systems (Bindeman et al., 1998; Coote and Shane, 2016). The majority of the plagioclase phenocrysts analyzed in this study are more enriched in MgO than the modeled melt compositions for a crystallization temperature of ~1099°C, which is based on the plagioclase geothermometer described above (Fig. 6a). This implies that the plagioclase in the PUBs analyzed in this study could not have crystallized from the basaltic host melt (MgO of 6.38%–8.87% (wt), average of 7.77% (wt)) and would require a more mafic melt (MgO of up to 14% (wt)). This agrees with the major element modeling (Ca/Na) of the plagioclase cores, but does not agree with the equilibrium between the rims and the melt suggested by the Ca/Na ratios (Fig. 5).

    Figure  6.  Compositional plots of An versus MgO (a) and FeO/MgO (b) for the plagioclase. The curves represent plagioclase equilibrium compositions based on the KMg values from Bindeman et al. (1998) and a temperature of 1099℃ obtained from the plagioclase geothermometer. The average crystallization temperature of the phenocryst rims is 1099°C. The following melt parameters were used: (1) MgO=14%, the upper limit of the MgO content of the matrix melt according to a simulation based on the model of Bindeman et al. (1998); (2) MgO=7.77%, the average whole-rock composition of the aphyric basalts in the same segment from the literature, which is representative of the matrix.

    The rimward increase in Mg and decrease in An appear to be similar to the trends described in previous diffusion studies (Costa et al., 2003; Moore et al., 2014). When minerals crystallize due to large degrees of undercooling and the diffusion in the melt cannot keep pace with the crystal formation, it is possible for late–stage rapid disequilibrium crystallization to produce a boundary layer melt enriched in incompatible (Fe, Mg) elements (Ginibre et al., 2002; Coote et al., 2018). This could explain the elevated MgO contents of the plagioclase rims and the groundmass plagioclase relative to the equilibrium values (Fig. 6a). The internal disequilibrium in the MgO contents can be explained by the mixing of primitive (MgO=14% (wt)) and more evolved magmas (MgO=7.77%(wt)).

    Additionally, in plagioclase, the post–crystallization diffusion of Fe is slower than that of Mg (Costa et al., 2003). Thus, the highest FeO/MgO ratios occur in the outermost parts of the rims and in the groundmass plagioclase (Fig. 6b) due to the fact that FeO diffuses into the plagioclase more slowly than MgO, and thus, more FeO accumulates in the boundary layer during a short residence time. However, it is worth noting that since the FeO content of the plagioclase depends on both the melt composition and the oxygen fugacity, it is difficult to assess the plagioclase–melt equilibrium using only the FeO content (Coote et al., 2018). Therefore, disequilibrium diffusion within a short time period can result in elevated Mg contents compared with the equilibrium values and the highest FeO/MgO ratios occurring in the plagioclase rims (Coote and Shane, 2016; Moore et al., 2014). Whereas the discrepancies in the plagioclase–melt equilibrium inferred from the major elements (Ca/Na) and the Mg contents are likely an artifact of disequilibrium diffusion (Coote and Shane, 2016).

    In the SWIR, the crystallization temperature and composition of the plagioclase cores are distinctly different from those of the rims (Table 1), which indicates that the plagioclase cores and rims have different thermal histories.

    The plagioclase cores are usually uniform and exhibit oscillatory zoning. The oscillatory zoning results in small-scale compositional variations, which suggests a regime of near–constant intensive parameters (pressure, temperature) (Landi et al., 2004; Shcherbakov et al., 2011). These characteristics suggest the cores crystallized from a stable environment and do not have complex crystallization histories. Besides, the presence of cores in plagioclase that are too primitive to be in equilibrium with the host magma indicates that they crystallized from a more primitive region in the plumbing system and were picked up by a more evolved melt later. Thus, we propose that the plagioclase cores grew in a stable mush zone where the temperature was high and constant, and were later entrained into a more evolved melt.

    The thin plagioclase rims (normally < 150 μm) have lower crystallization temperatures and lower An values than the cores, and their major elements (Ca/Na) are in equilibrium with the host magma. These characteristics demonstrate that they crystallized from the host melt, which is more evolved than the magma from which the cores crystalized. Similarly, the plagioclase microphenocrysts and microlites are also in equilibrium (Ca/Na) values with the host magma, which suggests that they crystallized from the host magma as well. Compared with fast to intermediate spreading ridges, there are generally no stable magma chambers and the volume of melt may be low under ultraslow mid–ocean ridges (Dick et al., 2003). Thus, the transport of magma into cooler regions of the reservoir would be expected to result in abrupt, strong undercooling of the magma. Due to these large degrees of rapid cooling, the plagioclase phenocrysts in SWIR PUBs generally have thin rims. The plagioclase rims of the phenocrysts exhibit major element (Fe, Mg) enrichment because these elements cannot reach equilibrium when diffusing from the host magma into the plagioclase phenocrysts. The MgO contents of the outermost rims of the plagioclase can be used to calculate the maximum time between the incorporation of the plagioclase into the host melt and quenching on the seafloor (Costa et al., 2003; Moore et al., 2014). We used Eq. (8) in Costa et al. (2003) to determine the Mg diffusion coefficient of the plagioclase. The maximum residence time of the plagioclases in the host melt was nearly 5–8 d. The close temporal relationship between the evolved magma replenishment and the eruption suggests that replenishment plays an important role in driving the eruption, which has also been suggested for other MORB eruptions on slow and intermediate ridges (Costa et al., 2010). Overall, our favored model is that replenishment by an evolved melt under the SWIR ridges (51°E) drives the eruption over a short period of time.

    (1) The plagioclase cores with high An values have higher crystallization temperatures (1 273±18)°C than the rims (1 099±10)°C. The range of crystallization temperatures for the microphenocrysts and microlites in the groundmass is similar to that of the phenocryst rims.

    (2) The compositions of the plagioclase cores from the SWIR indicate that they did not form in the host magma, but xenocrysts are crystallized from a more mafic melt composition. Whereas the plagioclase rims and the microphenocrysts and microlites in the groundmass are in equilibrium with the host basaltic melts.

    (3) The disequilibrium MgO contents and the higher FeO/MgO ratios of the rims of the plagioclase phenocrysts reflect shorter magmatic residence time periods than would be resulting from equilibrium diffusion.

    (4) An evolved melt replenished the magma under the SWIR ridges (51°E), driving the eruption over a short period of time.

    We are grateful to Yin-Jia Jin and two anonymous reviewers for their careful editing and constructive comments, which improved the manuscript. We also thank the crew and scientists involved in the R/V Dayang Yihao Cruise DY115–21.

  • Altschul S F, Madden T L, Schäffer A A, et al. 1997. Gapped BLAST and PSI-BLAST: a new generation of protein database search programs. Nucleic Acids Res, 25(17): 3389-3402
    Amo T, Atomi H, Imanaka T. 2003. Biochemical properties and regulated gene expression of the superoxide dismutase from the facultatively aerobic hyperthermophile Pyrobaculum calidifontis. J Bacteriol, 185(21): 6340-6347
    An S S, Kim Y M. 1997. Purification and characterization of a manganese-containing superoxide dismutase from a carboxydobacterium, Pseudomonas carboxydohydrogena. Mol Cells, 7(6): 730-737
    Angelova M, Dolashka-Angelova P, Ivanova E, et al. 2001. A novel glycosylated Cu/Zn-containing superoxide dismutase: production and potential therapeutic effect. Microbiology, 147(Pt6): 1641- 1650
    Asada K, Yoshikawa K, Takahashi M, et al. 1975. Superoxide dismutases from a blue-green alga, Plectonema boryanum. J Biol Chem, 250(8): 2801-2807
    Bannister J V, Bannister W H, Rotilio G. 1987. Aspects of the structure, function and applications of superoxide dismutase. CRC Crit Rev Biochem, 22(2): 111-180
    Beauchamp C, Fridovich I. 1971. Superoxide dismutase: improved assays and an assay applicable to acrylamide gels. Anal Biochem, 44(1): 276-287
    Bostwick D G, Alexander E E, Singh R, et al. 2000. Antioxidant enzyme expression and reactive oxygen species damage in prostatic intraepithelial neoplasia and cancer. Cancer, 89(1): 123-134
    Boyadzhieva I P, Atanasova M, Emanuilova E. 2010. A novel, thermostable manganese-containing superoxide dismutase from Bacillus licheniformis. Biotechnol Lett, 32(12): 1893-1896
    Bradford M M. 1976. A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. Anal Biochem, 72(1-2): 248-254
    Cullen J J, Weydert C, Hinkhouse M M, et al. 2003. The role of manganese superoxide dismutase in the growth of pancreatic adenocarcinoma. Cancer Res, 63(6): 1297-1303
    Dello Russo A, Rullo R, Nitti G, et al. 1997. Iron superoxide dismutase from the archaeon Sulfolobus solfataricus: average hydrophobicity and amino acid weight are involved in the adaptation of proteins to extreme environments. Biochim Biophys Acta, 1343(1): 23-30
    Emerit J, Samuel D, Pavio N. 2006. Cu-Zn super oxide dismutase as a potential antifibrotic drug for hepatitis C related fibrosis. Biomed Pharmacother, 60(1): 1-4
    Fridovich I. 1975. Superoxide dismutases. Annu Rev Biochem, 44(1): 147-159
    Guo Fangxian, E Shijin, Liu Shouan, et al. 2008. Purification and characterization of a thermostable MnSOD from the thermophilic fungus Chaetomium thermophilum. Mycologia, 100(3): 375-380
    Hakamada Y, Koike K, Kobayashi T, et al. 1997. Purification and properties of mangano-superoxide dismutase from a strain of alkaliphilic Bacillus. Extremophiles, 1(2): 74-78
    Hassan H M. 1989. Microbial superoxide dismutases. Adv Genet, 26: 65-97
    He Yongzhi, Fan Keqiang, Jia Cuijuan, et al. 2007. Characterization of a hyperthermostable Fe-superoxide dismutase from hot spring. Appl Microbiol Biotechnol, 75(2): 367-376
    Kardinahl S, Anemüller S, Schäfer G. 2000. The hyper-thermostable Fesuperoxide dismutase from the archaeon Acidianus ambivalens: characterization, recombinant expression, crystallization and effects of metal exchange. Biol Chem, 381(11): 1089-1101
    Klenk H P, Schleper C, Schwass V, et al. 1993. Nucleotide sequence, transcription and phylogeny of the gene encoding the superoxide dismutase of Sulfolobus acidocaldarius. Biochim Biophys Acta, 1174(1): 95-98
    Knapp S, Kardinahl S, Hellgren N, et al. 1999. Refined crystal structure of a superoxide dismutase from the hyperthermophilic archaeon Sulfolobus acidocaldarius at 2.2 Å resolution. J Mol Biol, 285(2): 689-702
    Laemmli U K. 1970. Cleavage of structural proteins during the assembly of the head of bacteriophage T4. Nature, 227(5259): 680-685
    Lancaster V L, LoBrutto R, Selvaraj F M, et al. 2004. A cambialistic superoxide dismutase in the thermophilic photosynthetic bacterium Chloroflexus aurantiacus. J Bacteriol, 186(11): 3408-3414
    Lee H J, Kwon H W, Koh J U, et al. 2010. An efficient method for the expression and reconstitution of thermostable Mn/Fe superoxide dismutase from Aeropyrum pernix K1. J Microbiol Biotechnol, 20(4): 727-731
    Li Duochuan, Gao Jing, Li Yaling, et al. 2005. A thermostable manganese-containing superoxide dismutase from the thermophilic fungus Thermomyces lanuginosus. Extremophiles, 9(1): 1-6
    Lim J H, Yu Y G, Choi I G, et al. 1997. Cloning and expression of superoxide dismutase from Aquifex pyrophilus, a hyperthermophilic bacterium. FEBS Lett, 406(1-2): 142-146
    Liu Bin, Wang Yiqian, Zhang Xiaobo. 2006. Characterization of a recombinant maltogenic amylase from deep sea thermophilic Bacillus sp. WPD616. Enzyme Microb Tech, 39(4): 805-810
    Liu Jianguo, Yin Mengmeng, Zhu Hu, et al. 2011. Purification and characterization of a hyperthermostable Mn-superoxide dismutase from Thermus thermophilus HB27. Extremophiles, 15(2): 221- 226
    Luisa Corvo M, Jorge J C, van't Hof R, et al. 2002. Superoxide dismutase entrapped in long-circulating liposomes: formulation design and therapeutic activity in rat adjuvant arthritis. Biochim Biophys Acta, 1564(1): 227-236
    Melov S, Ravenscroft J, Malik S, et al. 2000. Extension of life-span with superoxide dismutase/catalase mimetics. Science, 289(5484): 1567-1569
    McCord J M. 1976. Iron-and manganese-containing superoxide dismutases: structure, distribution, and evolutionary relationships. Adv Exp Med Biol, 74: 540-550
    McCord J M, Fridovich I. 1969. Superoxide dismutase. An enzymic function for erythrocuprein (hemocuprein). J Biol Chem, 244(22): 6049-6055
    McCord J M, Fridovich I. 1988. Superoxide dismutase: the first twenty years (1968-1988). Free Radic Biol Med, 5(5-6): 363-369
    Miroshnichenko M L, Bonch-Osmolovskaya E A. 2006. Recent developments in the thermophilic microbiology of deep-sea hydrothermal vents. Extremophiles, 10(2): 85-96
    Nishikawa M, Nagatomi H, Nishijima M, et al. 2001. Targeting superoxide dismutase to renal proximal tubule cells inhibits nephrotoxicity of cisplatin and increases the survival of cancer-bearing mice. Cancer Lett, 171(2): 133-138
    Oppenheimer C H, ZoBell C E. 1952. The growth and viability of sixtythree species of marine bacteria as influenced by hydrostatic pressure. J Mar Res, 11: 10-18
    Parker M W, Blake C C. 1988. Iron-and manganese-containing superoxide dismutases can be distinguished by analysis of their primary structures. FEBS Lett, 229(2): 377-382
    Raimondi S, Uccelletti D, Matteuzzi D, et al. 2008. Characterization of the superoxide dismutase SOD1 gene of Kluyveromyces marxianus L3 and improved production of SOD activity. Appl Microbiol Biotechnol, 77(6): 1269-1277
    Song Chongfu, Sheng Liangquan, Zhang Xiaobo. 2012. Preparation and characterization of a thermostable enzyme (Mn-SOD) immobilized on supermagnetic nanoparticles. Appl Microbiol Biotechnol, 96(1): 123-132
    Song Ningning, Zheng Yan, E Shijin, et al. 2009. Cloning, expression, and characterization of thermostable manganese superoxide dismutase from Thermoascus aurantiacus var. levisporus. J Microbiol, 47(1): 123-130
    Stewart R R C, Bewley J D. 1980. Lipid peroxidation associated with accelerated aging of soybean axes. Plant Physiol, 65(2): 245-248
    Ursby T, Adinolfi B S, Al-Karadaghi S, et al. 1999. Iron superoxide dismutase from the archaeon Sulfolobus solfataricus: analysis of structure and thermostability. J Mol Biol, 286(1): 189-205
    Valderas M W, Hart M E. 2001. Identification and characterization of a second superoxide dismutase gene (sodM) from Staphylococcus aureus. J Bacteriol, 183(11): 3399-3407
    Vieille C, Zeikus G J. 2001. Hyperthermophilic enzymes: sources, uses, and molecular mechanisms for thermostability. Microbiol Mol Biol Rev, 65(1): 1-43
    Vorauer-Uhl K, Fürnschlief E, Wagner A, et al. 2001. Topically applied liposome encapsulated superoxide dismutase reduces postburn wound size and edema formation. Eur J Pharm Sci, 14(1): 63-67
    Wang Xin, Yang Haijie, Ruan Lingwei, et al. 2008. Cloning and characterization of a thermostable superoxide dismutase from the thermophilic bacterium Rhodothermus sp. XMH10. J Ind Microbiol Biotechnol, 35(2): 133-139
    Weisiger R A, Fridovich I. 1973. Mitochondrial superoxide simutase. Site of synthesis and intramitochondrial localization. J Biol Chem, 248(13): 4793-4796
    Whittaker M M, Whittaker J W. 2000. Recombinant superoxide dismutase from a hyperthermophilic archaeon, Pyrobaculum aerophilium. J Biol Inorg Chem, 5(3): 402-408
    Wu Suijie, Liu Bin, Zhang Xiaobo. 2006. Characterization of a recombinant thermostable xylanase from deep-sea thermophilic Geobacillus sp. MT-1 in East Pacific. Appl Microbiol Biotechnol, 72(6): 1210-1216
    Yabe Y, Kobayashi N, Nishihashi T, et al. 2001. Prevention of neutrophil-mediated hepatic ischemia/reperfusion injury by superoxide dismutase and catalase derivatives. J Pharmacol Exp Ther, 298(3): 894-899
    Yamano S, Sako Y, Nomura N, et al. 1999. A cambialistic SOD in a strictly aerobic hyperthermophilic archaeon, Aeropyrum pernix. J Biochem, 126(6): 218-225
    Yoo H Y, Kim S S, Rho H M. 1999. Overexpression and simple purification of human superoxide dismutase (SOD1) in yeast and its resistance to oxidative stress. J Biotechnol, 68(1): 29-35
    Yost F J Jr, Fridovich I. 1973. An iron-containing superoxide dismutase from Escherichia coli. J Biol Chem, 248: 4905-4908
    Youn H D, Kim E J, Roe J H, et al. 1996. A novel nickel-containing superoxide dismutase from Streptomyces spp. Biochem J, 318(Pt3): 889-896
    Yu Jing, Yu Xiaomin, Liu Jianhua. 2004. A thermostable manganesecontaining superoxide dismutase from pathogen Chlamydia pneumoniae. FEBS Lett, 562(1-3): 22-26
    Yu Ping. 2007. A new approach to the production of the recombinant SOD protein by methylotrophic Pichia pastoris. Appl Microbiol Biotechnol, 74(1): 93-98
    Yunoki M, Kawauchi M, Ukita N, et al. 2003. Effects of lecithinized superoxide dismutase on neuronal cell loss in CA3 hippocampus after traumatic brain injury in rats. Surg Neurol, 59(3): 156-160
    Zhang Liqing, Guo Fangxian, Xian Hongquan, et al. 2011. Expression of a novel thermostable Cu, Zn-superoxide dismutase from Chaetomium thermophilum in Pichia pastoris and its antioxidant properties. Biotechnol Lett, 33(6): 1127-1132
    Zhang Yi, Wang Junzhi, Wu Yongjie, et al. 2002. Anti-inflammatory effect of recombinant human superoxide dismutase in rats and mice and its mechanism. Acta Pharmacol Sin, 23(5): 439-444
    Zhong Weixiong, Oberley L W, Oberley T D, et al. 1997. Suppression of the malignant phenotype of human glioma cells by overexpression of manganese superoxide dismutase. Oncogene, 14(4): 481-490
    Zhu Yanbing, Li Hebin, Zhang Xuqin, et al. 2011. Characterization of a thermostable manganese-containing superoxide dismutase from inshore hot spring thermophile Thermus sp. JM1. Acta Oceanol Sin, 30(6): 95-103
    Zou Yuanyuan, Yang Ling, Liu Guoxian, et al. 2011. Identification of three superoxide dismutase genes from a Geobacillus sp.. Protein J, 30(1): 66-71
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