High Mn2 Research Articles

Solid‐state NMR spectra of paramagnetic silica‐based materials: observation of 29Si and 27Al nuclei in the first coordination spheres of manganese ions

Some silica-based solids, prepared by the sol/gel method in the presence of high Mn2+ concentrations, have been characterized by the 29Si, 27Al MAS NMR spectra and 29Si T1 measurements. The single-pulse 29Si and 27Al MAS NMR spectra have shown broad spinning sideband patterns that are interpreted in terms of anisotropic bulky magnetic susceptibility (BMS) and dipole-field effects. In the absence of paramagnetic isotropic shifts, the 29Si and 27Al nuclei observed in the single-pulse NMR spectra have been assigned to nuclei remote from paramagnetic centers. It has been demonstrated that the 29Si and 27Al nuclei, which are in the vicinity of the manganese ions, can be detected by the Hahn-echo MAS NMR experiments at different carrier frequencies.

Magnetic Study of Two Isotypic Manganese Chloro‐Sulfides: MnSbS2Cl and the New Compound MnBiS2Cl.

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Mn 2+ -Dependent Regulation of Multiple Genes in Streptococcus pneumoniae through PsaR and the Resultant Impact on Virulence

The concentration of Mn2+ is 1,000-fold higher in secretions than it is at internal sites of the body, making it a potential signal by which bacteria can sense a shift from a mucosal environment to a more invasive site. PsaR, a metal-dependent regulator in Streptococcus pneumoniae, was found to negatively affect the transcription of psaBCA, pcpA, rrgA, rrgB, rrgC, srtBCD, and rlrA in the presence of Mn2+. psaBCA encode an ABC-type transporter for Mn2+. pcpA, rrgA, rrgB, and rrgC encode several outer surface proteins. srtBCD encode a cluster of sortase enzymes, and rlrA encodes a transcriptional regulator. Steady-state RNA levels are high under low Mn2+ concentrations in the wild-type strain and are elevated under both high and low Mn2+ concentrations in a psaR mutant strain. RlrA is an activator of rrgA, rrgB, rrgC, and srtBCD (D. Hava and A. Camilli, Mol. Microbiol. 45:1389-1406, 2002), suggesting that PsaR may indirectly control these genes through rlrA, while PsaR-dependent repression of psaBCA, pcpA, and rlrA transcription is direct. The impact of Mn2+-dependent regulation on virulence was further examined in mouse models of pneumonia and nasopharyngeal carriage. The abilities of DeltapsaR, pcpA, and DeltapsaR DeltapcpA mutant strains to colonize the lung were reduced compared to those of the wild type, confirming that both PcpA-mediated gene regulation and PsaR-mediated gene regulation are required for full virulence in the establishment of pneumonia. Neither PcpA nor PsaR was found to be required for colonization of the nasopharynx in a carriage model. This is the first demonstration of Mn2+ acting as a signal for the expression of virulence factors within different host sites.

The Influence of Mn2+on DNA structure in the presence of Na+ions: A Raman spectroscopic study

The influence of Mn2+ions on the structure of natural calf thymus DNA was studied by Raman spectroscopy. Measurements were done at room temperature and pH 6.2±0.2, in the presence of the physiological concentration of 150 mM Na+ions, and in the presence of Mn2+concentrations that varied between 0 and 600 mM. No condensation of DNA was observed at any of the Mn2+concentrations. At 5 mM Mn2+and 150 mM Na+no significant influence of Mn2+ions on the DNA structure can be observed. Compared with our previous results obtained at 10 mM Na+ions, binding of Mn2+ions to charged phosphate groups and to DNA bases is inhibited in the presence of 150 mM Na+ions. DNA backbone conformational changes were not observed in the whole concentration range of Mn2+ions as judging from the Raman spectra. No evidence for DNA melting was identified. A high Mn2+affinity for binding to guanine N7 and possibly, in a much lesser extent, to adenine have been found.

Involvement of tannase in the acquisition of manganese from tannin-rich medium by tannase-producing Lactobacillus plantarum

The role of tannase produced by Lactobacillus plantarum in tannin-rich medium was demonstrated by using strains with known tannase activities. After 20 h incubation in aerobic tannin-rich medium, the growth of L. plantarum KOG 19 with low activity was lower than that of L. plantarum ATCC 14917T with high activity by approximately 1.5 orders of magnitude, whereas the growth of KOG 19 in aerobic medium without tannin was almost the same as that of ATCC 14917T. The reduced growth of KOG 19 in aerobic tannin-rich medium was recovered by the addition of exogenous tannase or Mn2+ to the medium. A similar result was obtained in the comparison of growth levels between high and low tannase activity groups in aerobic tannin-rich medium, while no significant differences were observed between the groups in anaerobic tannin-rich medium. Furthermore, the growth of the low tannase activity group in aerobic tannin-rich medium was restored by the addition of catalase. These results suggest that tannase production allows L. plantarum strains to acquire and accumulate Mn2+ which is otherwise chelated by tannins, and then high intracellular Mn2+ compensates for the lack of the enzyme superoxide dismutase in L. plantarum resulting in resistance to oxygen toxicity under aerobic conditions.

Micro-x-ray absorption near-edge structure imaging for detecting metallic Mn in GaN

In this study, we report the application of a synchrotron radiation microprobe to the analysis of Mn valencies in GaN. X-ray absorption near-edge structure (XANES) images taken around MnK-edge in fluorescence detection mode reveal the concentration of oxidation states of Mn centers. By fitting the XANES curve for each point of the image, the distributions of the Mn0, Mn2+, and Mn3+ oxidation states are obtained. At low Mn concentrations, there is a homogeneous mixture of Mn2+ and Mn3+ centers, while at high Mn content strong spatial-dependent Mn0 and Mn2+ distributions characterize the XANES maps. In a supplementary way with respect to Mn2+, the Mn0 pattern suggests the presence of specific cluster-like features, indicating surface segregation of metallic Mn centers.

A Novel Isoform of the Secretory Pathway Ca2+,Mn2+-ATPase, hSPCA2, Has Unusual Properties and Is Expressed in the Brain

Unlike lower eukaryotes, mammalian genomes have a second gene, ATP2C2, encoding a putative member of the family of secretory pathway Ca2+,Mn(2+)-ATPases, SPCA2. Human SPCA2 shares 64% amino acid identity with the protein defective in Hailey Hailey disease, hSPCA1. We show that human SPCA2 (hSPCA2) has a more limited tissue distribution than hSPCA1, with prominent protein expression in brain and testis. In primary neuronal cells, endogenous SPCA2 has a highly punctate distribution that overlaps with vesicles derived from the trans-Golgi network and is thus different from the compact perinuclear distribution of hSPCA1 seen in keratinocytes and nonpolarized cells. Heterologous expression in a yeast strain lacking endogenous Ca2+ pumps reveals further functional differences from hSPCA1. Although the Mn(2+)-specific phenotype of hSPCA2 is similar to that of hSPCA1, Ca2+ ions are transported with much poorer affinity, resulting in only weak complementation of Ca(2+)-specific yeast phenotypes. These observations suggest that SPCA2 may have a more specialized role in mammalian cells, possibly in cellular detoxification of Mn2+ ions, similar to that in yeast. We point to the close links between manganese neurotoxicity and Parkinsonism that would predict an important physiological role for SPCA2 in the brain.

The effect of manganese(II) on DNA structure: electronic and vibrational circular dichroism studies

The interaction of DNA with Mn2+ was studied in absorbance and optical activity in the electronic and vibrational regions. Based on the data, several stages of the interaction were identified. Con formational transition towards the C-form of DNA was observed in solution at the molar ratio Mn2+/DNA-phosphates between 0.1 and 1.5. The exact ratio depended on the ionic strength and increased with increasing NaCl concentration. Although manganese interacted with the phosphates and bases of DNA at higher metal concentrations, it is unlikely that direct chelation occurred. A model for the interaction between manganese ions and DNA mediated by water is suggested destabilizing the double helix and partially breaking the hydrogen bonds between the base pairs. At high Mn2+ concentrations DNA aggregation was observed.

Two-metal ion mechanism of RNA cleavage by HIV RNase H and mechanism-based design of selective HIV RNase H inhibitors

Human immunodeficiency virus (HIV) RNase H activity is essential for the synthesis of viral DNA by HIV reverse transcriptase (HIV-RT). RNA cleavage by RNase H requires the presence of divalent metal ions, but the role of metal ions in the mechanism of RNA cleavage has not been resolved. We measured HIV RNase H activity associated with HIV-RT protein in the presence of different concentrations of either Mg2+, Mn2+, Co2+ or a combination of these divalent metal ions. Polymerase-independent HIV RNase H was similar to or more active with Mn2+ and Co2+ compared with Mg2+. Activation of RNase H by these metal ions followed sigmoidal dose-response curves suggesting cooperative metal ion binding. Titration of Mg2+-bound HIV RNase H with Mn2+ or Co2+ ions generated bell-shaped activity dose-response curves. Higher activity could be achieved through simultaneous binding of more than one divalent metal ion at intermediate Mn2+ and Co2+ concentrations, and complete replacement of Mg2+ occurred at higher Mn2+ or Co2+ concentrations. These results are consistent with a two-metal ion mechanism of RNA cleavage as previously suggested for a number of polymerase-associated nucleases. In contrast, the structurally highly homologous RNase HI from Escherichia coli is most strongly activated by Mg2+, is significantly inhibited by submillimolar concentrations of Mn2+ and most probably cleaves RNA via a one-metal ion mechanism. Based on this difference in active site structure, a series of small molecule N-hydroxyimides was identified with significant enzyme inhibitory potency and selectivity for HIV RNase H.

Effect of Free Manganese Activity on Yield and Uptake of Micronutrient Cations by Barley and Oat grown in Chelator-buffered Nutrient Solution

The uptake of micronutrient cations in relation to varying activities of Mn2+ was studied for barley (Hordeum vulgare L. var. Thule) and oat (Avena sativa L. var. Biri) grown in chelator buffered nutrient solution. Free activities of Mn2+ were calculated by using the chemical speciation programme GEOCHEM-PC. Manganese deficient conditions induced elevated concentrations of Zn and Fe in shoots of both species. The corresponding antagonistic relationship between Mn and Cu could only be seen in barley. The observed antagonistic relationships were only valid as long as the plant growth was limited by Mn deficiency. The Mn concentration in both plant species increased significantly with increasing Mn2+ activity in the nutrient solution. The concentration of Mn in the shoots of oat was higher than for barley except under severe Mn deficiency where it was found equal for both species. Manganese was accumulated in the roots of barley at high Mn2+ activity. The different shoot concentrations of Mn between barley and oat are therefore attributed to the extent of Mn translocation from roots to shoots. Manganese deficiency induced a significant increase in the shoot to root ratio in both species.

Optical and Surface Structural Properties of Mn2+-Doped ZnSe Nanoparticles

Mn2+-doped ZnSe nanoparticles were synthesized from molecular cluster precursors. Four ZnSe nanoparticle samples, one with low Mn2+ concentration (A), one with an intermediate Mn2+ concentration (B), one with a high Mn2+ concentration (C), and one with no Mn2+, were prepared and characterized using UV−vis, luminescence, electron spin resonance (ESR), and X-ray absorption fine structure (XAFS) techniques. The sample with no Mn2+ had a sharp ZnSe band edge emission peak and a quantum yield of ∼2%. The samples with Mn2+ had a significant decrease in band edge emission. Sample A had no Mn2+ 4T1 → 6A1 emission but showed some ZnSe band edge emission and trap state emission. Sample B had Mn2+ 4T1 → 6A1 emission and a further reduction in ZnSe band edge emission and trap state emission. Sample C showed an increase in the Mn2+ 4T1 → 6A1 emission, a dramatic increase in trap state emission, and essentially no ZnSe band edge emission. The overall emission from all four samples was quenched with time. To better unde...

Manganês e germinação de esporos de fungos micorrízicos arbusculares in vitro

A alta disponibilidade de íons metálicos no solo, dentre eles o Mn2+, pode inibir os fungos micorrízicos arbusculares (FMA), retardando a germinação dos esporos e, conseqüentemente, a formação de micorriza, o que reduz a eficiência simbiótica. O objetivo deste trabalho foi avaliar o efeito do íon Mn2+ sobre a germinação de esporos de seis espécies de FMA dos gêneros Glomus, Acaulospora, Gigaspora e Scutellospora em experimento in vitro Em substrato constituído por areia lavada, adicionaram-se 15; 30 e 75 mg kg-1 de Mn2+, na forma de MnCl2.4H2O, mantendo-se um controle sem adição de Mn. Acondicionaram-se os esporos em envelopes membranosos, introduzindo-os entre duas camadas de areia com diferentes níveis de Mn, em placas de Petri. Após 30 dias de incubação, avaliou-se a germinação dos esporos. Houve decréscimo médio de 32, 49 e 75 % na germinação dos esporos, à medida que se aumentaram as doses de Mn, em comparação ao controle. A germinação de esporos do gênero Glomus foi totalmente inibida na maior dose. O gênero Acaulospora sofreu decréscimos de até 50 % já na dose de 15 mg kg-1 de Mn2+, enquanto os gêneros Scutellospora e Gigaspora apresentaram os maiores índices de germinação de esporos, tendo havido tolerância no caso de Gigaspora, mesmo na maior dose de Mn2+, e estímulo à germinação no caso de Scutellospora até à dose 30 mg kg-1 de Mn2+.

Manganese‐enhanced MRI of mouse heart during changes in inotropy

Recently the dual properties of manganese ion (Mn2+) as an MRI contrast agent and a calcium analogue to enter excitable cells has been used to mark specific cells in brain and as a potential intracellular cardiac contrast agent. Here the hypothesis that in vivo manganese-enhanced MRI (MEMRI) can detect changes in inotropy in the mouse heart has been tested. T1-weighted images were acquired every minute during an experimental time course of 75 min. Varying doses of Mn2+ (3.3–14.0 nmoles/min/g BW) were infused during control and altered inotropy with dobutamine (positive inotropy due to increased calcium influx) and the calcium channel blocker diltiazem (negative inotropy). Infusion of MnCl2 led to a significant increase in signal enhancement in mouse heart that saturated above 3.3 ± 0.1 nmoles/min/g BW Mn2+ infusion. At the highest Mn2+ dose infused there was a 41–47% increase in signal intensity with no alteration in cardiac function as measured by MRI-determined ejection fractions. Dobutamine increased both the steady-state level of enhancement and the rate of MRI signal enhancement. Diltiazem decreased both the steady-state level of enhancement and the rate of MRI signal enhancement. These results are consistent with the model that Mn2+-induced enhancement of cardiac signal is indicative of the rate of calcium influx into the heart. Thus, the simultaneous measurement of global function and calcium influx using MEMRI may provide a useful method of evaluating in vivo responses to inotropic therapy. Magn Reson Med 46:884–890, 2001. Published 2001 Wiley-Liss, Inc.

Factors Influencing the Luminescence Quantum Efficiency of Nanocrystalline ZnS:Mn2+

The influence of the Mn2+ concentration, the nature of the passivating polymer and UV irradiation on the luminescence efficiency of nanocrystalline ZnS:Mn2+ is investigated. With increasing Mn2+ concentration the quantum efficiency increases, until it reaches a stable value. No drastic decrease in quantum efficiency is observed at high Mn2+ concentrations, as was found earlier for both bulk and nanocrystalline ZnS:Mn2+. By altering the passivating polymer many other factors influencing the quantum efficiency are changed, such as the particle diameter and Mn content. This makes it hard to study the influence of the passivating behaviour of the various polymers on the quantum efficiency. The UV enhancement of the quantum efficiency of the various samples could be explained by two mechanisms: UV curing of the passivating polymer and passivation by a photo-oxidation of the surface of the nanoparticles during the irradiation. The highest quantum efficiency obtained was about 15%.

Effects of Mn 2+ and NH + 4 concentrations on laccase and manganese peroxidase production and Amaranth decoloration by Trametes versicolor

Extracellular lignin peroxidase (LiP) was not detected during decoloration of the azo dye, Amaranth, by Trametes versicolor. Approximately twice as much laccase and manganese peroxidase (MnP) was produced by decolorizing cultures compared to when no dye was added. At a low Mn2+ concentration (3 M), N-limited (1.2 mM NH4+) cultures decolorized eight successive additions of Amaranth with no visible sorption to the mycelial biomass. At higher Mn2+ concentrations (200 M), production of MnP increased and that of laccase decreased, but the rate or number of successive Amaranth decolorations was unaffected. There was always a 6-h to 8-h lag prior to decoloration of the first aliquot of Amaranth, regardless of MnP and laccase concentrations. Although nitrogen-rich (12 mM NH4+) cultures at an initial concentration of 200 M Mn2+ produced high laccase and MnP levels, only three additions of Amaranth were decolorized, and substantial mycelial sorption of the dye occurred. While the results did not preclude roles for MnP and laccase, extracellular MnP and laccase alone were insufficient for decoloration. The cell-free supernatant did not decolorize Amaranth, but the mycelial biomass separated from the whole broth and resuspended in fresh medium did. This indicates the involvement of a mycelial-bound, lignolytic enzyme or a H2O2-generating mechanism in the cell wall. Nitrogen limitation was required for the expression of this activity.

Activation/Attenuation Model for RNase H: A ONE-METAL MECHANISM WITH SECOND-METAL INHIBITION

Ribonucleases H (RNases H) comprise a family of metal-dependent enzymes that catalyze the hydrolysis of the 3'-O---P bond of RNA in RNA.DNA hybrids. The mechanism by which RNases H use active-site metal(s) for catalysis is unclear. Based upon the seemingly contradictory structural observations of one divalent metal bound to Escherichia coli RNase HI and two divalent metals bound to the HIV RNase H domain, two models explaining RNase H metal dependence have been proposed: a one-metal mechanism and a two-metal mechanism. In this paper, we show that the Mn2+-dependent activity of E. coli RNase HI is not consistent with either of these mechanisms. RNase H activity in the presence of Mn2+ is complex, with activation and inhibition of the enzyme at low and high Mn2+ concentrations, respectively. Mutations at Asp-134 result in a partial loss of this inhibition, with little effect on activation. Neutralization of His-124 by mutation to Ala results in an enzyme with a significantly decreased specific activity and an absolute loss of Mn2+ inhibition. Inhibition by high Mn2+ concentrations is shown to be due to a reduction in kcat; this attenuation has a critical dependence on the presence of His-124. Based upon these results, we propose an "activation/attenuation" model explaining the metal dependence of RNase H activity where one metal is required for enzyme activation and binding of a second metal is inhibitory.

Kinetics of Mn3+–Oxalate Formation and Decay in Reactions Catalyzed by Manganese Peroxidase ofCeriporiopsis subvermispora

The kinetics of Mn3+–oxalate formation and decay were investigated in reactions catalyzed by manganese peroxidase (MnP) from the basiomyceteCeriporiopsis subvermisporain the absence of externally added hydrogen peroxide. A characteristic lag observed in the formation of this complex was shortened by glyoxylate or catalytic amounts of Mn3+or hydrogen peroxide. MnP titers had a minor effect on this lag and did not influence the decay rate of the complex. In contrast, Mn2+and oxalate drastically affected maximal concentrations of the Mn3+–oxalate complex formed, the decay of which was accelerated at high Mn2+levels. The highest concentration of complex was obtained at pH 4.0, whereas an inverse relationship was found between the pH of the reaction and the decay rate of the complex with MnP present. In the absence of MnP, the best fit for the decay kinetics of the complex gave an order of 3/2 at concentrations in the range of 30–100 μM, with akobs= 0.012 min−1M−0.5at pH 4.0. The rate constant increases at lower pH values and decreases at high oxalate concentrations. The physiological relevance of these findings is discussed.

Differential effects of the reticulospinal system on locomotion in lamprey.

Specific effects of stimulating different parts of the reticulospinal (RS) system on the spinal locomotor pattern are described in lamprey. In the in vitro brain stem and spinal cord preparation, microstimulation in different areas of the reticular formation was performed by ejecting a small amount of -glutamate from a micropipette. These areas were distributed over the four reticular nuclei of the brain stem: the mesencephalic reticular nucleus (MRN) and the anterior, middle and posterior rhombencephalic reticular nuclei (ARRN, MRRN, and PRRN, respectively). To prevent synaptic spread of excitation within the brain stem, the synaptic transmission was blocked by using a low Ca2+, high Mn2+ physiological saline in the brain stem pool. "Fictive" locomotion was evoked by applying N-methyl--aspartate (NMDA) to the spinal cord. Rhythmical discharges of motoneurons were recorded bilaterally in the midbody area, from the ventral roots that had been subdivided in dorsal and ventral branches, supplying the dorsal and ventral part of the myotome, respectively. Two major effects of brain stem stimulation were elicited: a change in the frequency of the locomotory rhythm and an induction of asymmetry (left/right, dorsal/ventral) in the segmental motor output. Approximately 50% of the stimulated sites evoked a change in locomotor frequency. In the PRRN almost all effective sites evoked an increase in frequency (10-50%). In the other nuclei, increase and decrease (10-30%) were observed equally frequently. Most of the stimulated sites (50-80%) in any reticular nucleus evoked asymmetry in the segmental motor output. Distortion of the segmental output symmetry was classified into eight categories by comparing the intensity of locomotor bursts in the dorsal and ventral branches of the two ventral roots, ipsilateral and contralateral to the stimulated side. These categories differed in the direction of the body flexion, which would be evoked during normal swimming: ipsilateral (I), contralateral (C), dorsal (D), ventral (V), ipsilateral and dorsal (ID), ipsilateral and ventral (IV), contralateral and dorsal (CD), and contralateral and ventral (CV). The different categories were not equally represented in each nucleus and across the nuclei. The most pronounced categories for each nucleus were as follow. In MRN: I (33%); ARRN: C (44%); MRRN: rostral part, I (36%) and caudal part, CV (42%); and PRRN: rostral part, I (40%) and caudal part, IV (35%). Other categories were also present but less common in each nucleus. To examine if the effects of brain stem stimulation were uniform along the spinal cord, recordings were performed from distal parts of the cord. Stimulation of a given point in the brain stem produced similar pattern of effects in 59% of cases and different patterns in 41% of cases. The main conclusion of the present study is that the proportion of RS neurons with different influences on the spinal locomotor network differs significantly among different parts of the reticular formation of the lamprey. The specificity of RS influences may represent a basis for modifications of the segmental locomotor output necessary for the control of equilibrium and steering during locomotion.

Manganese uptake and toxicity in magnesium-supplemented and unsupplemented Saccharomyces cerevisiae.

The magnesium content of Saccharomyces cerevisiae was found to vary by up to fivefold at differing stages of batch growth and during growth in the presence of differing magnesium concentrations. Excess Mg was primarily sequestered in vacuoles. Mn(2+)-uptake experiments revealed that Mg-enriched cells had a markedly reduced capacity for Mn2+ accumulation. For example, after 6 h incubation in the presence of 50 microM Mn2+, Mn levels were approximately twofold higher in cells previously grown in unsupplemented medium than in those from Mg-supplemented medium. These differences were further accentuated at higher Mn2+ concentrations and were not attributable to altered cell-surface charge or altered cell-surface Mn2+ binding. Cellular Mg status also influenced Mn toxicity towards S. cerevisiae. During exposure to 5 mM Mn2+, 50% reductions in the viability of cells with initial Mg contents of approximately 1400 and 2700 nmol (10(9) cells)-1 occurred after approximately 1.6 h and 3.6 h respectively. In cells containing 3300 nmol Mg (10(9) cells)-1, more than 75% viability was still maintained after 7 h incubation with 5 mM Mn2+. It is concluded that Mn2+ uptake and toxicity in S. cerevisiae are strongly influenced by intracellular Mg, possibly through Mg-dependent regulation of divalent-cation transport activity.

Lignin-degrading activity and ligninolytic enzymes of different white-rot fungi: effects of manganese and malonate

Ten species of white-rot fungi, mainly belonging to the family Polyporaceae (Basidiomycotina), were studied in terms of their ability to degrade14C-ring labelled synthetic lignin and secrete ligninolytic enzymes in liquid cultures under varying growth conditions. Lignin mineralization by the fungi in an air atmosphere did not exceed 14% within 29 days. Different responses to the elevated Mn2+concentration and the addition of a manganese chelator (sodium malonate) were observed among various fungal species. This could be related with the utilization of either lignin peroxidase (LiP) or manganese peroxidase (MnP) for lignin depolymerization, i.e., some fungi apparently had an LiP-dominating ligninolytic system and others an MnP-dominating ligninolytic system. The LiP isoforms were purified from Trametes gibbosa and Trametes trogii. Isoelectric focusing of purified ligninolytic enzymes revealed the expression of numerous MnP isoforms in Trametes gibbosa, Trametes hirsuta, Trametes trogii, and Abortiporus biennis grown under a high (50-fold) Mn2+level (120 μM) with the addition of the chelator. In addition, two to three laccase isoforms were detected. Key words: white-rot fungi, lignin degradation, lignin peroxidase, manganese peroxidase, manganese, malonate.