Contribution of biomineralization during growth of polymetallic nodules and ferromanganese crusts from the Pacific Ocean

doi:10.1007/s11706-009-0033-0

Front Mater Sci Chin

0, Vol.

Issue () : 109-123 https://doi.org/10.1007/s11706-009-0033-0

RESEARCH ARTICLE

Contribution of biomineralization during growth of polymetallic nodules and ferromanganese crusts from the Pacific Ocean

Xiao-hong WANG¹(

), Guan Lu¹, Werner E. G. MüLLER²(

)

1. National Research Center for Geoanalysis, 26 Baiwanzhuang Dajie, Beijing 100037, China; 2. Institut für Physiologische Chemie, Abteilung Angewandte Molekularbiologie, Universit?t, Duesbergweg 6, D-55099 Mainz, Germany

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Abstract

The ocean hosts inorganic raw materials to a magnitude, which surpasses the resources of these materials available on land. Those mineral resources include industrial minerals, metalliferous oxides, hydrothermal metalliferous sulfides, and dissolved minerals. Hence, a significant source of minerals for sustainable recovery in the future may be ocean waters. Among of those mineral resources, there are two kinds of very important minerals which are consolidated on the seabeds of ocean basins in polymetallic nodules and on the surface of seamounts in polymetallic crusts. Until now, the (bio)-chemical processes that result in the formation of metal deposits in the form of nodules or crusts are not understood. In the present review, we concentrate on the (potential) biogenic origin of nodule and crust formation.

We studied polymetallic/ferromanganese nodules that had been collected from the Clarion-Clipperton Zone in the Eastern Pacific Ocean, by high-resolution scanning electron microscopy (HR-SEM) to search for microorganisms. The nodules are made up of small-sized micronodules, 100 to 450 μm in size, which are bound/glued together by an interstitial whitish material. In these micronodules, dense accumulations of microorganisms/bacteria can be visualized that display only two morphotypes: (i) round-shaped cocci and (ii) elongated rods. The microorganisms are decorated on their surfaces with S-layers, which are indicative for bacteria. Moreover, the data suggest that these S-layers are the crystallization seeds for the mineralization process. We conclude that the mineral material of the nodule has a biogenic origin and propose consequently the view that mineralization in nodules is caused by biologically controlled mineralization processes.

In a second series of investigations, first evidence for a biogenic origin of ferromanganese crusts formation is given. Crusts were obtained from the Magellan seamounts and analyzed for their chemical composition using the EDX technique. Again, special emphasis had been put on the (potential) biogenic origin of the mineral deposition in these ferromanganese crusts. We could demonstrate by HR-SEM that, in those deposits, vast amounts of coccoliths (calcareous unicellular algae) exist. Surprisingly, the coccoliths are composed of Mn besides Ca and C, as analyzed by EDX. This result could be further substantiated by EDX mappings. We propose that initiation of crust formation involves the dissolution of calcite from the coccoliths, resulting in an oxidation of Mn²⁺ to Mn⁴⁺ and subsequent precipitation of Mn⁴⁺O₂. Following this scheme, it can be assumed that crust formation may serve as an example for a biologically induced mineralization process.

Keywords polymetallic nodules ferromanganese crusts bacteria coccolithophores microorganisms biomineralization biogenic materials

Corresponding Author(s): WANG Xiao-hong,Email:wxh0408@hotmail.com (Prof. Dr. X.H. WANG); MüLLER Werner E. G.,Email:wmueller@uni-mainz.de (Prof. Dr. W.E.G. MüLLER)

Issue Date: 05 June 2009

Cite this article:

Xiao-hong WANG,Guan Lu,Werner E. G. MüLLER. Contribution of biomineralization during growth of polymetallic nodules and ferromanganese crusts from the Pacific Ocean[J]. Front Mater Sci Chin, 0, (): 109-123.

URL:

https://academic.hep.com.cn/foms/EN/10.1007/s11706-009-0033-0
https://academic.hep.com.cn/foms/EN/Y0/V/I/109

Fig.1 First descriptions of nodules and crusts: A close up of polymetallic nodules, collected in 1891 during the Challenger Expedition [] in the Coral Sea close to New Guinea. The sizes of those nodules vary around 5 cm. During the German Deep Sea Expedition “Valdivia” (1898-1899), large deposits (here, 8-cm large samples) of phosphorite crusts have been described around Cape of Good Hope (Africa).

Fig.2 Scheme proposing processes, generating the formation of hy-Fe/Mn-crusts and the polymetallic nodules in the deep sea. Modification, based on Halbach [] and Koschinsky et al. []. It is outlined that the formation of the hy-Fe/Mn-crusts proceeds in the mixing zone between the upper oxygen-minimum zone and the lower oxygen-rich deep water. The latter bottom water originates from the two sources, the Pacific Deep Water (PDW) and the Antarctic Bottom Water (AABW). The morphology of the crusts displays a anthracite layer, which is usually secondarily phosphatized (in red); a middle separating speck layer (yellow); and an upper layer (brownish). The crust contacts directly the Basaltic substrate of the seamount. Two types of polymetallic nodules exist. According to their genesis, they are subdivided in diagenetically formed nodules (originated by early diagenesis) and hydrogenetic nodules (hydrogenesis). Both of them grow in deeper zones than the crusts; the diagenetic nodules (di-nodules) grow in the pelagic sediments, whereas the hydrogenetic nodules (hy-nodules) are formed on the surface of the sediments (se).

Fig.3 Nodule morphology. Nodules were collected from the Clarion-Clipperton Zone in the Pacific Ocean: Outer appearance of a nodule, with its smooth surface texture. View to the major axis of a nodule, grouped to the elongated morphotypes. The rubber net (n) stabilizes the nodule. Polished section through a nodule, which comprises at least four macrolamellae (1 to 4). A cross breakage of a nodule displays the lamellar structure (la) with a pale gray submetallic luster. The outer lamella is dark brown to black, suggesting a composition of birnessite (bi). The individual lamellae are separated by a zone that comprises a dendritic, ornamentous pattern. This pattern is assembled and composed of single blackish drops, termed micronodules (mn). Those micronodules are dominant in the surface lamellae. (Digital light microscopy, performed with a VHX-600 Digital Microscope from KEYENCE.) The micro-nodules (mn) become well distinguishable after breaking the nodules. (HR-SEM analyses using a Gemini Leo 1530 high-resolution field emission scanning electron microscope.) Formation of hydrogenetic nodules (hy-nodules); scheme. Initially, minute clay/sand aggregates form the substrate for the adhesion of bacteria. Mediated by the microorganisms, soluble metallic ions (Mn,Fe, Cuand Co) are metabolized by oxidation or salt formation into insoluble biogenic minerals. Subsequently, the surfaces of the micro-organisms function as seed templates and allow a progressing mineralization through non-biogenic processes. Subsequently, micronodules are formed by accretion and inclusion of bacteria in non-biogenic mineral material. Finally, the micronodules assemble to nodules, via rotating movements on the sea floor, under simultaneous inclusion of inorganic material.

Fig.4 Microorganisms, abundantly present in micronodules, cocci, and rods. HR-SEM analysis: within the micronodules, both cocci (co) and rods (ro) are seen. Individual coccus (co); its smooth surface is covered with small-sized platelets, presumably consisting of oxides. Frequently, cocci arranged in bead-like chains are found that are known from the genus Streptococcus. Where present, rods (ro) are associated with cocci (co) in the micronodules. One rod chain (ro(c)) is shown. The rods are aligned in chains (rod-chain; ro(c)), providing division septa (s) within the linearly oriented rods or rod chain. Occasionally, organisms, reminiscent of coccolithophores, are seen.

Fig.5 EDX analyses of defined regions within the nodules (SEM-EDX analysis). A square with an edge length of 10 μm has been used for the EDX determinations. A region in the outer zone of a nodule, comprising micronodules (mn). Those micronodules are rich in bacteria/microorganisms.The areas from which the EDX analyses have been performed, refer to Fig. 5(B). SEM/EDX analyses from the different regions in a nodule, comprising micronodules. Analyses have been performed on a micro-nodule ((B)-a), within a nest of micronodules ((B)-b), or in an area, surrounding the micronodules (interstitial material) ((B)-c). These locations are also marked in Fig. 5(A).

Fig.6 Microbial assemblages in nodules. : in the outer region of nodules (exolithobiontic colonization), in their microcanals, filamentous (likely to be) bacteria/actinobacteria (fb) exist, which comprises a smooth surface. In addition to the filamentous bacteria (fb), spherical to helical bacteria/microorganisms (sb) are found to expose central deep openings. These taxa do not show any indication for a mineralization process. : Visualization of the biofilms within the nodules by HR-SEM (endolithobiontic colonization). Organization of cone-like microorganisms (operationally abbreviated with b (bacteria)) existing on biofilm platforms. The matrix support (m) for the microorganisms as well as the holes (h), previously harboring them, are marked. Hexagonal microorganisms (b) arranged in an (almost) perfect honeycomb pattern. All of these stone/pillar-like microorganisms are covered by brick-like mineral deposits. EDX spectra from selected areas within a biofilm, formed by cone-like microorganisms (b). Overview of a biofilm (bf). The area (approximately 1 μm) outside of the biofilm that had been analyzed by EDX is marked (I-a). Higher magnification of a region, occupied with cone-like microorganisms (b), assembled at a biofilm (bf) matrix. Those EDX analyses have been performed from a region of a bacterium (I-b) and the spacing between them (I-c). Corresponding EDX spectra from selected areas within the biofilm in (G) and (H). Spectrum from an outside region of the biofilm ((I)-a); spectrum taken from a cone-like microorganism ((I)-b) and spectrum from a biofilm region adjacent to the microorganisms ((I)-c).

Fig.7 S-layers existing on bacteria. Cone-like structures (operationally abbreviated here with: b; standing for bacteria) on a plane area within nodules. The phalanx of cones is arranged in parallel lines on the surface of the plane structure, representing a previous biofilm (bf). The individual cones have a size of 0.8 to 1.0 μm and are regularly arranged in a pattern with an interspacing of 1.0 to 1.5 μm. Horizontal view onto a subsurface layer, comprising the cone-like structures (b). Identification of S-layers, crystalline structures, on the surfaces of the cones. Each individual bacterium (b; size: 800×300 nm) is decorated at its outward directed surface, with 20 to 25 pillar-shaped protrusions (p); (size: 75×45 nm), representing S-layers. The arrangement of the protrusions follows basically an oblique to square pattern. HR-SEM images. Schematic representation of the process of mineral deposition in nodules by endolithic bacteria. It is highly conceivable that these microorganisms exist in different morphotypes; in the top layer (i) of the mineral deposits, the bacteria with their S-layers are seen. These structures facilitate the initial stages of crystallization. (ii) With progressing of mineral deposition, the microorganisms become covered with the efflorescenting mineral deposits, which have a brick-like morphology. (iii) The shape of the microorganisms turns to cone-like, and finally, (iv) the minerals at the lateral surfaces of the microorganisms fuse and acquire the same texture as the base substratum. It can be proposed that, during this biogenic mineralization in the nodules, the initially amorphous metal deposits undergo diagenesis to a semicrystalline state.

Fig.8 Proposed biogenic deposition of minerals on bacteria, from which the growth of nodules originates (scheme). Ectoenzymes (E), proposed to exist on the surface of the microbes, oxidize Mn to Mn that are largely insoluble and form the seeds for the subsequent mineralization processes. In addition, it is proposed that these enzymes are arranged at the particles (nm-size), comprising the S-layer (surface layer) of the bacteria.

Fig.9 Morphology of the crust sample. A vertical breakage of the sample reveals a distinct zonation; a lower layer (ll) and a middle layer (“speck layer” (sl)), which separates the lower layer from the upper layer (ul). The basis (ba) of the crust, which contacts Basaltic rock, is characterized by the red encrusts. A direct view to the basis of the crust discloses the spotty appearance with the red encrusts. A plane cross-section through the crust; upper layer. Individual convex bags are stacked upon each other in a direction (see direction of arrows), which runs parallel to the surface (su).

Fig.10 Elemental mapping of a section through the surface layers of the stacked individual convex bags (cb). The directions of growth of the bags are indicated with arrows. Electronic image (EI). Scanning for Mn concentration (Mn) and Scanning for Ca concentration. The relative concentrations increase with a change in color from dark/blue to yellow to red.

Fig.11 Well-preserved assemblages from coccoliths are present in the crust; HR-SEM. A broken crust displaying islands of regions that are rich in Mn and contain abundantly coccospheres (cos). Higher magnification of such an area displaying individual coccolithophores (col). Those coccolithophore assemblies are composed of calcified scales, the coccoliths (coc). X-ray energy spectrum (area 20 μm) of an area, rich in coccoliths.

Fig.12 EDX maps of transverse fracture through a crust, rich in coccoliths. HR-SEM image analyzing a single coccolith. EDX map for calcium (Ca) and manganese (Mn).

Fig.13 Free energy relationships between CaCO dissolution (from coccoliths) via the intermediate Mn()CO to Mn()O (in crusts). These reactions are driven by pH shifts and concentration differences.

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