• Title/Summary/Keyword: ClC channel

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Bicuculline Methiodide (BMI) Induces Membrane Depolarization of The Trigeminal Subnucleus Caudalis Substantia Gelatinosa Neuron in Mice Via Non-$GABA_A$ Receptor-Mediated Action

  • Yin, Hua;Park, Seon-Ah;Choi, Soon-Jeong;Bhattarai, Janardhan P.;Park, Soo-Joung;Suh, Bong-Jik;Han, Seong-Kyu
    • International Journal of Oral Biology
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    • v.33 no.4
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    • pp.217-221
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    • 2008
  • Bicuculline is one of the most commonly used $GABA_A$ receptor antagonists in electrophysiological research. Because of its poor water solubility, bicuculline quaternary ammonium salts such as bicuculline methiodide (BMI) and bicuculline methbromide are preferred. However, a number of studies have shown that BMI has non-$GABA_A$ receptor-mediated effects. The substantia gelatinosa (SG) of the trigeminal subnucleus caudalis (Vc) is implicated in the processing of nociceptive signaling. In this study, we investigated whether BMI has non-GABA receptor-mediated activity in Vc SG neurons using a whole cell patch clamp technique. SG neurons were depolarized by application of BMI ($20{\mu}M$) using a high $Cl^-$ pipette solution. GABA ($30-100{\mu}M$) also induced membrane depolarization of SG neuron. Although BMI is known to be a $GABA_A$ receptor antagonist, GABA-induced membrane depolarization was enhanced by co-application with BMI. However, free base bicuculline (fBIC) and picrotoxin (PTX), a $GABA_A$ and $GABA_C$ receptor antagonist, blocked the GABA-induced response. Furthermore, BMI-induced membrane depolarization persisted in the presence of PTX or an antagonist cocktail consisting of tetrodotoxin ($Na^+$ channel blocker), AP-5 (NMDA receptor antagonist), CNQX (non-NMDA receptor antagonist), and strychnine (glycine receptor antagonist). Thus BMI induces membrane depolarization by directly acting on postsynaptic Vc SG neurons in a manner which is independent of $GABA_A$ receptors. These results suggest that other unknown mechanisms may be involved in BMI-induced membrane depolarization.

Medical Image Compression Using JPEG International Standard (JPEG 표준안을 이용한 의료 영상 압축)

  • Ahn, Chang-Beom;Han, Sang-Woo;Kim, Il-Yoen
    • Proceedings of the KIEE Conference
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    • 1993.07a
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    • pp.504-506
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    • 1993
  • The Joint Photographic Experts Group (JPEG) standard was proposed by the International Standardization Organization (ISO/SC 29/WG 10) and the CCITT SG VIII as an international standard for digital continuous-tone still image compression. The JPEG standard has been widely accepted in electronic imaging, computer graphics, and multi-media applications, however, due to the lossy character of the JPEG compression its application in the field of medical imaging has been limited. In this paper, the JPEG standard was applied to a series of head sections of magnetic resonance (MR) images (256 gray levels, $256{\times}256$ size) and its performance was investigated. For this purpose, DCT-based sequential mode of the JPEG standard was implemented using the CL550 compression chip and progressive and lossless coding was implemented by software without additional hardware. From the experiment, it appears that the compression ratio of about 10 to 20 was obtained for the MR images without noticeable distortion. It is also noted that the error signal between the reconstructed image by the JPEG and the original image was nearly random noise without causing any special-pattern-related artifact. Although the coding efficiency of the progressive and hierarchical coding is identical to that of the sequential coding in compression ratio and SNR, it has useful features In fast search of patient Image from huge image data base and in remote diagnosis through slow public communication channel.

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Effects of Cyclobuxine E on Two Distinct Types of Potassium-Activated Calcium Channels in an Intestinal Smooth Muscle (흰쥐 장관에 있어 칼륨에 의해 활성화되는 칼슘 채널에 대한 Cyclobuxine E의 영향)

  • Lee, Jong-Hwoa;Kwon, Jun-Tack;Cho, Byung-Heon;Choi, Kyu-Hong;Kim, Yu-Jae;Kim, Jong-Bae;Kim, Chun-Sook;Cha, Young-Deog;Kim, Young-Suk
    • The Korean Journal of Pharmacology
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    • v.25 no.1
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    • pp.53-58
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    • 1989
  • For several years, we investigated the pharmacological action of several substances isolated from Buxus microphylla var koreana Nakai, which had been used as folk remedies of malaria and venereal disease. Cyclobuxine $D(C_{25}H_{42}ON_2)$, a steroidal alkaloid, exerted an antiinflammatory action, hypotensive and bradycardic effects in rats. In the present study, we isolated alkaloid from the acetone-insoluble fraction of the strong bases of this plants. This alkaloid $(C_{25}H_{38}ON_2)$ was identified as a steroidal alkaloid contained a cyclopropane ring by physical and chemical methods. It is a derivative of cyclobuxine D and named cyclobuxine E. We examined the effect of cyclobuxine E on the contractile response induced by acetylcholine and two distinct types of potassium-activated calcium channels in an intestinal smooth muscle of the rat. Cyclobuxine E inhibited significantly the Ach-induced contraction. The isolated longitudinal muscle from the rat duodenum was immersed calcium-depleted potassium depolarizing solution. Ten minutes after, 1.8 mM $CaCl_2$ was added to muscle bath and elicited a biphasic increase in muscle tension. Cyclobuxine E produced an appreciable inhibition of both components of the mechanical response. In addition, Cyclobuxine E introduced at a point when the tonic response had reached its maximum level, caused the muscle to exhibit a rapid loss of tension. Based on these experimental results, we proposed the possibility that the inhibitory action of cyclobuxine E on the isolated rat duodenum may be due to inhibiting the transmembrane fluxes of calcium ion in potassium-activated calcium channels.

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Fabrication of Polycrystalline Si Films by Silicide-Enhanced Rapid Thermal Annealing and Their Application to Thin Film Transistors (Silicide-Enhanced Rapid Thermal Annealing을 이용한 다결정 Si 박막의 제조 및 다결정 Si 박막 트랜지스터에의 응용)

  • Kim, Jone Soo;Moon, Sun Hong;Yang, Yong Ho;Kang, Sung Mo;Ahn, Byung Tae
    • Korean Journal of Materials Research
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    • v.24 no.9
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    • pp.443-450
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    • 2014
  • Amorphous (a-Si) films were epitaxially crystallized on a very thin large-grained poly-Si seed layer by a silicide-enhanced rapid thermal annealing (SERTA) process. The poly-Si seed layer contained a small amount of nickel silicide which can enhance crystallization of the upper layer of the a-Si film at lower temperature. A 5-nm thick poly-Si seed layer was then prepared by the crystallization of an a-Si film using the vapor-induced crystallization process in a $NiCl_2$ environment. After removing surface oxide on the seed layer, a 45-nm thick a-Si film was deposited on the poly-Si seed layer by hot-wire chemical vapor deposition at $200^{\circ}C$. The epitaxial crystallization of the top a-Si layer was performed by the rapid thermal annealing (RTA) process at $730^{\circ}C$ for 5 min in Ar as an ambient atmosphere. Considering the needle-like grains as well as the crystallization temperature of the top layer as produced by the SERTA process, it was thought that the top a-Si layer was epitaxially crystallized with the help of $NiSi_2$ precipitates that originated from the poly-Si seed layer. The crystallinity of the SERTA processed poly-Si thin films was better than the other crystallization process, due to the high-temperature RTA process. The Ni concentration in the poly-Si film fabricated by the SERTA process was reduced to $1{\times}10^{18}cm^{-3}$. The maximum field-effect mobility and substrate swing of the p-channel poly-Si thin-film transistors (TFTs) using the poly-Si film prepared by the SERTA process were $85cm^2/V{\cdot}s$ and 1.23 V/decade at $V_{ds}=-3V$, respectively. The off current was little increased under reverse bias from $1.0{\times}10^{-11}$ A. Our results showed that the SERTA process is a promising technology for high quality poly-Si film, which enables the fabrication of high mobility TFTs. In addition, it is expected that poly-Si TFTs with low leakage current can be fabricated with more precise experiments.

Expression and Purification of the Phosphatase-like Domain of a Voltage-Sensing Phosphatase, Ci-VSP (막 전위 감지 탈인산화 효소, Ci-VSP의 유사 탈인산화 효소 도메인의 발현과 정제)

  • Kim, Sung-Jae;Kim, Hae-Min;Choi, Hoon;Kim, Young-Jun
    • Journal of Life Science
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    • v.21 no.7
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    • pp.1032-1038
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    • 2011
  • Recently identified Ciona intestinalis voltage sensor-containing phosphatase (Ci-VSP) consists of an ion channel-like transmembrane domain (VSD) and a phosphatase-like domain. Ci-VSP senses the change of membrane potential by its VSD and works as a phosphoinositide phosphatase by its phosphatase domain. In this study, we present the construction of His-tagged phosphatase-like domain of Ci-VSP, its recombinant expression and purification, and its enzymatic activity behavior in order to examine the biochemical behavior of phosphatase domain of Ci-VSP without interference. We found that Ci-VSP(248-576)-His can be eluted with an elution buffer containing 25 mM NaCl and 100 mM imidazole during His-tag purification. In addition, we found the proper measurement condition for kinetics study of Ci-VSP(248-576)-His against p-nitrophenyl phosphate (pNPP). We measured the kinetic constant of Ci-VSP(248-576)-His at $37^{\circ}C$, pH 5.0 or 5.5, under 30 min of reaction time, and less than $2.0\;{\mu}g$ of protein amount. With these conditions, we acquired that Ci-VSP(248-576)-His has $K_m$ of $354{\pm}0.143\;{\mu}M$, $V_{max}$ of $0.0607{\pm}0.0137\;{\mu}mol$/min/mg and $k_{cat}$ of $0.359{\pm}0.009751\;min^{-1}$ for pNPP dephosphorylation. Therefore, we produced a pure form of Ci-VSP(248-576)-His, and this showed a higher activity against pNPP. This purified protein will provide the road to a structural investigation on an interesting protein, Ci-VSP.

The Inhibitory Effects of Hydrogen Sulfide on Pacemaker Activity of Interstitial Cells of Cajal from Mouse Small Intestine

  • Parajuli, Shankar Prasad;Choi, Seok;Lee, Jun;Kim, Young-Dae;Park, Chan-Guk;Kim, Man-Yoo;Kim, Hyun-Il;Yeum, Cheol-Ho;Jun, Jae-Yeoul
    • The Korean Journal of Physiology and Pharmacology
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    • v.14 no.2
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    • pp.83-89
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    • 2010
  • In this study, we studied whether hydrogen sulfide ($H_2S$) has an effect on the pacemaker activity of interstitial cells of Cajal (ICC), in the small intestine of mice. The actions of $H_2S$ on pacemaker activity were investigated using whole-cell patch-clamp technique, intracellular $Ca^{2+}$ analysis at $30^{\circ}C$ and RT-PCR in cultured mouse intestinal ICC. Exogenously applied sodium hydrogen sulfide (NaHS), a donor of hydrogen sulfide, caused a slight tonic inward current on pacemaker activity in ICC at low concentrations (50 and $100{\mu}m$), but at high concentration ($500{\mu}m$ and 1 mM) it seemed to cause light tonic inward currents and then inhibited pacemaker amplitude and pacemaker frequency, and also an increase in the resting currents in the outward direction. Glibenclamide or other potassium channel blockers (TEA, $BaCl_2$, apamin or 4-aminopydirine) did not have an effect on NaHS-induced action in ICC. The exogenous application of carbonilcyanide p-triflouromethoxyphenylhydrazone (FCCP) and thapsigargin also inhibited the pacemaker activity of ICC as NaHS. Also, we found NaHS inhibited the spontaneous intracellular $Ca^{2+}$ ($[Ca^{2+}]_i$) oscillations in cultured ICC. In doing an RT-PCR experiment, we found that ICC enriched population lacked mRNA for both CSE and CBS, but was prominently detected in unsorted muscle. In conclusion, $H_2S$ inhibited the pacemaker activity of ICC by modulating intracellular $Ca^{2+}$. These results can serve as evidence of the physiological action of $H_2S$ as acting on the ICC in gastrointestinal (GI) motility.

Novel Glycolipoproteins from Ginseng

  • Pyo, Mi-Kyung;Choi, Sun-Hye;Hwang, Sung-Hee;Shin, Tae-Joon;Lee, Byung-Hwan;Lee, Sang-Mok;Lim, Yoong-Ho;Kim, Dong-Hyun;Nah, Seung-Yeol
    • Journal of Ginseng Research
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    • v.35 no.1
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    • pp.92-103
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    • 2011
  • Ginseng has been used as a general tonic agent to invigorate human body. In the present study, we isolated novel glycolipoproteins from ginseng that activate $Ca^{2+}$-activated $Cl^-$ channel (CaCC) in Xenopus oocytes and transiently increase intracellular free $Ca^{2+}$ concentration ($[Ca^{2+}]_i$) in mouse Ehrlich ascites tumor cells. We named the active ingredients as gintonin. Gintonin exists in at least six different forms. The native molecular weight of gintonin is about 67 kDa but its apparent molecular weight is about 13 kDa, indicating that gintonin might be a pentamer. Gintonin is rich in hydrophobic amino acids. Its main carbohydrates are glucose and glucosamine. Its lipid components are linoleic, palmitic, oleic, and stearic acids. Gintonin actions were blocked by U73122, a phospholipase C inhibitor, 2-aminoethxydiphenyl borate, an inositol 1,4,5-trisphosphate receptor antagonist, or bis (o-aminophenoxy) ethane-N,N,N0,N0-tetracetic acid acetoxymethyl ester, a membrane permeable $Ca^{2+}$ chelator. In the present study, we for the first time isolated novel gintonin and showed the signaling pathways on gintonin-mediated CaCC activations and transient increase of $[Ca^{2+}]_i$. Since $[Ca^{2+}]_i$ as a second messenger plays a pivotal role in the regulation of diverse $Ca^{2+}$-dependent intracellular signal pathways, gintonin-mediated regulations of $[Ca^{2+}]_i$ might contribute to biological actions of ginseng.

Sesquiterpenoids Bioconversion Analysis by Wood Rot Fungi

  • Lee, Su-Yeon;Ryu, Sun-Hwa;Choi, In-Gyu;Kim, Myungkil
    • 한국균학회소식:학술대회논문집
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    • 2016.05a
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    • pp.19-20
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    • 2016
  • Sesquiterpenoids are defined as $C_{15}$ compounds derived from farnesyl pyrophosphate (FPP), and their complex structures are found in the tissue of many diverse plants (Degenhardt et al. 2009). FPP's long chain length and additional double bond enables its conversion to a huge range of mono-, di-, and tri-cyclic structures. A number of cyclic sesquiterpenes with alcohol, aldehyde, and ketone derivatives have key biological and medicinal properties (Fraga 1999). Fungi, such as the wood-rotting Polyporus brumalis, are excellent sources of pharmaceutically interesting natural products such as sesquiterpenoids. In this study, we investigated the biosynthesis of P. brumalis sesquiterpenoids on modified medium. Fungal suspensions of 11 white rot species were inoculated in modified medium containing $C_6H_{12}O_6$, $C_4H_{12}N_2O_6$, $KH_2PO_4$, $MgSO_4$, and $CaCl_2$ for 20 days. Cultivation was stopped by solvent extraction via separation of the mycelium. The metabolites were identified as follows: propionic acid (1), mevalonic acid lactone (2), ${\beta}$-eudesmane (3), and ${\beta}$-eudesmol (4), respectively (Figure 1). The main peaks of ${\beta}$-eudesmane and ${\beta}$-eudesmol, which were indicative of sesquiterpene structures, were consistently detected for 5, 7, 12, and 15 days These results demonstrated the existence of terpene metabolism in the mycelium of P. brumalis. Polyporus spp. are known to generate flavor components such as methyl 2,4-dihydroxy-3,6-dimethyl benzoate; 2-hydroxy-4-methoxy-6-methyl benzoic acid; 3-hydroxy-5-methyl phenol; and 3-methoxy-2,5-dimethyl phenol in submerged cultures (Hoffmann and Esser 1978). Drimanes of sesquiterpenes were reported as metabolites from P. arcularius and shown to exhibit antimicrobial activity against Gram-positive bacteria such as Staphylococcus aureus (Fleck et al. 1996). The main metabolites of P. brumalis, ${\beta}$-Eudesmol and ${\beta}$-eudesmane, were categorized as eudesmane-type sesquiterpene structures. The eudesmane skeleton could be biosynthesized from FPP-derived IPP, and approximately 1,000 structures have been identified in plants as essential oils. The biosynthesis of eudesmol from P. brumalis may thus be an important tool for the production of useful natural compounds as presumed from its identified potent bioactivity in plants. Essential oils comprising eudesmane-type sesquiterpenoids have been previously and extensively researched (Wu et al. 2006). ${\beta}$-Eudesmol is a well-known and important eudesmane alcohol with an anticholinergic effect in the vascular endothelium (Tsuneki et al. 2005). Additionally, recent studies demonstrated that ${\beta}$-eudesmol acts as a channel blocker for nicotinic acetylcholine receptors at the neuromuscular junction, and it can inhibit angiogenesis in vitro and in vivo by blocking the mitogen-activated protein kinase (MAPK) signaling pathway (Seo et al. 2011). Variation of nutrients was conducted to determine an optimum condition for the biosynthesis of sesquiterpenes by P. brumalis. Genes encoding terpene synthases, which are crucial to the terpene synthesis pathway, generally respond to environmental factors such as pH, temperature, and available nutrients (Hoffmeister and Keller 2007, Yu and Keller 2005). Calvo et al. described the effect of major nutrients, carbon and nitrogen, on the synthesis of secondary metabolites (Calvo et al. 2002). P. brumalis did not prefer to synthesize sesquiterpenes under all growth conditions. Results of differences in metabolites observed in P. brumalis grown in PDB and modified medium highlighted the potential effect inorganic sources such as $C_4H_{12}N_2O_6$, $KH_2PO_4$, $MgSO_4$, and $CaCl_2$ on sesquiterpene synthesis. ${\beta}$-eudesmol was apparent during cultivation except for when P. brumalis was grown on $MgSO_4$-free medium. These results demonstrated that $MgSO_4$ can specifically control the biosynthesis of ${\beta}$-eudesmol. Magnesium has been reported as a cofactor that binds to sesquiterpene synthase (Agger et al. 2008). Specifically, the $Mg^{2+}$ ions bind to two conserved metal-binding motifs. These metal ions complex to the substrate pyrophosphate, thereby promoting the ionization of the leaving groups of FPP and resulting in the generation of a highly reactive allylic cation. Effect of magnesium source on the sesquiterpene biosynthesis was also identified via analysis of the concentration of total carbohydrates. Our current study offered further insight that fungal sesquiterpene biosynthesis can be controlled by nutrients. To profile the metabolites of P. brumalis, the cultures were extracted based on the growth curve. Despite metabolites produced during mycelia growth, there was difficulty in detecting significant changes in metabolite production, especially those at low concentrations. These compounds may be of interest in understanding their synthetic mechanisms in P. brumalis. The synthesis of terpene compounds began during the growth phase at day 9. Sesquiterpene synthesis occurred after growth was complete. At day 9, drimenol, farnesol, and mevalonic lactone (or mevalonic acid lactone) were identified. Mevalonic acid lactone is the precursor of the mevalonic pathway, and particularly, it is a precursor for a number of biologically important lipids, including cholesterol hormones (Buckley et al. 2002). Farnesol is the precursor of sesquiterpenoids. Drimenol compounds, bi-cyclic-sesquiterpene alcohols, can be synthesized from trans-trans farnesol via cyclization and rearrangement (Polovinka et al. 1994). They have also been identified in the basidiomycota Lentinus lepideus as secondary metabolites. After 12 days in the growth phase, ${\beta}$-elemene caryophyllene, ${\delta}$-cadiene, and eudesmane were detected with ${\beta}$-eudesmol. The data showed the synthesis of sesquiterpene hydrocarbons with bi-cyclic structures. These compounds can be synthesized from FPP by cyclization. Cyclic terpenoids are synthesized through the formation of a carbon skeleton from linear precursors by terpene cyclase, which is followed by chemical modification by oxidation, reduction, methylation, etc. Sesquiterpene cyclase is a key branch-point enzyme that catalyzes the complex intermolecular cyclization of the linear prenyl diphosphate into cyclic hydrocarbons (Toyomasu et al. 2007). After 20 days in stationary phase, the oxygenated structures eudesmol, elemol, and caryophyllene oxide were detected. Thus, after growth, sesquiterpenes were identified. Per these results, we showed that terpene metabolism in wood-rotting fungi occurs in the stationary phase. We also showed that such metabolism can be controlled by magnesium supplementation in the growth medium. In conclusion, we identified P. brumalis as a wood-rotting fungus that can produce sesquiterpenes. To mechanistically understand eudesmane-type sesquiterpene biosynthesis in P. brumalis, further research into the genes regulating the dynamics of such biosynthesis is warranted.

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Sulfide Chimney from the Cleft Segment, Juan de Fuca Ridge: Mineralogy and Fluid Inclusion (Juan de Fuca 해령 Cleft Segment에서 회수된 황화물 침니: 광물조성 및 유체포유물)

  • 윤성택;허철호;소칠섭;염승준;이경용
    • Economic and Environmental Geology
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    • v.35 no.3
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    • pp.203-210
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    • 2002
  • In order to elucidate the growth mechanism of sulfide chimney formed as a result of seafloor hydrothermal mineralization, we carried out the mineralogical and fluid inclusion studies on the inactive, sulfide- and silica-rich chimney which has been recovered from a hydrothermal field in the Cleft segment of the Juan de Fuca Ridge. According to previous studies, many active and inactive vents are present in the Cleft segment. The sulfide- and silica-rich chimney is composed of amorphous silica, pyrite, sphalerite and wurtzite with minor amounts of chalcopyrite and marcasite. The interior part of the chimney is highly porous and represents a flow channel. Open spaces within chimneys are typically coated with colloform layers of amorphous silica. The FeS content of Zn-sulfides varies widely from 13.9 to 34.3 mole% with Fe-rich core and Fe-poor rims. This variation possibly reflects the change of physicochemical characteristics of hydrothermal fluids. Chemical and mineralogical compositions of the each growth zone are also varied, possibly due to a thermal gradient. Based on the microthermometric measurements of liquid-rich, two-phase inclusions in amorphous silica that was precipitated in the late stage of mineralization, minimum trapping temperatures are estimated to be about 1140 to 145$^{\circ}$C with the salinities between 3.2 and 4.8 wt.% NaCI equiv. Although the actual fluid temperatures of the vent are not available, this study suggests that the lowtemperature conditions were predominant during the mineralization in the hydrothermal field at Cleft segment. Comparing with the previously reported chimney types, the morphology, colloform texture, bulk chemistry, and a characteristic mineral assemblage (pyrite + marcasite + wurtzite + amorphous silica) of this chimney indicate that the chimney have been formed from a relatively low-temperature (<250$^{\circ}$C) hydrothermal fluid that was changed by sluggish fluid flow and conductive cooling.

Temporal and Spatial Variability of the Middle and Lower Tropospheric Temperatures from MSU and ECMWF (MSU와 ECMWF에서 유도된 중간 및 하부 대류권 온도의 시 ${\cdot}$ 공간 변동)

  • Yoo, Jung-Moon;Lee, Eun-Joo
    • Journal of the Korean earth science society
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    • v.21 no.5
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    • pp.503-524
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    • 2000
  • Intercomparisons between four kinds of data have been done to estimate the accuracy of satellite observations and model reanalysis for middle and lower tropospheric thermal state over regional oceans. The data include the Microwave Sounding Units (MSU) Channel 2 (Ch2) brightness temperatures of NOAA satellites and the vertically weighted corresponding temperature of ECMWF GCM (1980-93). The satellite data for midtropospheric temperatures are MSU2 (1980-98) in nadir direction and SC2 (1980-97) in multiple scans, and for lower tropospheric temperature SC2R (1980-97). MSU2 was derived in this study while SC2 and SC2R were described in Spencer and Christy (1992a, 1992b). Temporal correlations between the above data were high (r${\ge}$0.90) in the middle and high latitudes, but low(r${\sim}$0.65) over the low latitude and more convective regions. Their values with SC2R which included the noises due to hydrometeors and surface emission were conspicuously low. The reanalysis shows higher correlation with SC2 than with MSU2 partially because of the hydrometeors screening. SC2R in monthly climatological anomalies was more sensitive to surface thermal condition in northern hemisphere than MSU2 or SC2. The first EOF mode for the monthly mean data of MSU and ECMWF shows annual cycle over most regions except the tropics. The mode in MSU2 over the Pacific suggests the east-west dipole due to the Walker circulation, but this tendency is not clear in other data. In the first and second modes for the Ch2 anomalies over most regions, the MSU and ECMWF data commonly indicate interannual variability due to El Ni${\tilde{n}$o and La Ni${\tilde{n}$a. The substantial disagreement between observations and model reanalysis occurs over the equatorial upwelling region of the western Pacific, suggesting uncertainties in the model parameterization of atmosphere-ocean interaction.

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