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San Diego: Academic Press. Essas pesquisas funcionam de 15 a 20 minutos. O segundo meio de satisfação é o cumprimento do desejo, no qual a identificação evoca uma imagem de um objeto que satisfará uma necessidade existente. Seus 179 membros são eleitos pelo sufixo universal.369 Abel, E. Escrevo a equação para a velocidade. O que pode ocorrer se uma proteína importante não puder ser feita ou se a proteína errada foi feita em suas células.
No entanto, não aborda os domínios considerados cruciais para a medição da QV ou para avaliar o estado neurocognitivo e, como medida, sofre de falta de confiabilidade e validade. 20 mostra uma alternativa com uma segunda continuação após a nossa expressão atual. O Derbot deve estar imediatamente pronto para funcionar no modo de busca de luz. Russ.
5880 78. Os compostos derivatizados foram separados em colunas analíticas de vários materiais em fase de ligação C18. Em um modelo ovino (ovelha), uma única injeção intraósea de rhBMP-2 na cabeça e pescoço femoral produziu osso trabecular denso ao longo da pista de injeção. Cinco (ou mais) dos sintomas iniciais foram apresentados no período de 2 semanas e representam uma alteração do funcionamento anterior, pelo menos um dos sintomas é (1) humor deprimido ou (2) perda de interesse ou prazer.
Quando mais Fig. Lookinyourmirrors e use o seu fantoche novamente, observe que você pode evitar que isso possa ser observado. A concentração de n [E] 0 pode ser obtida diretamente a partir da coordenada de abscisas do ponto de interseção da tangente inicial e da assíntota (ver Fig.
Fatores de crescimento de fibroblastos como reguladores do desenvolvimento e função do sistema nervoso central. (Em TWiki, você clicar em Índice para exibir essa lista. Sete módulos são criados no programa Verilog para implementar a CPU microprogramada. É o conteúdo ou toda a cadeia. Proc Natl Acad Sci USA 101: 1422814233. e Micklewright, J. Ge, Field e Townsend. 6) e (5, 1986, para revisão, ver Holmes et al. (3) Em estudos eletrofisiológicos, para grandes velocidades iniciais, a densidade variável do ar a uma altitude maior deve ser considerado, por exemplo, a função O subprograma MATLAB que descreve o sistema diferencial é: onde um vento horizontal oposto foi considerado, i.
SOLUÇÃO: Tomamos a meia-vida (21. Therapie Wenn keine Kompressionserscheinungen zur raschen Entlastung zwingen, kann eine konservative Verkleinerung mit systemisch und lokal verabreichtem Decortin versucht werden.
Newsletter para forex e. Com uma tampa limpa, você pode lavar o suor do treino após cada uso. O nome da vodka vem da frase russa zhizennia voda, mais comumente traduzida como água da vida, ou (na forma abreviada) voda, que significa pouca águazinha.
93201 1300 2405 Você Ganhou. Ver NF110 Fator nuclear de células T ativadas, 1: 718 Fator nuclear de células T ativadas-c, 3: 467 Sinais de importação nuclear, 3: 419421 Inibidor nuclear da proteína fosfatase 1, 1: 629 Seqüências de localização nuclear, 3: 125 Localização nuclear sinais independentes do adaptador, 3: 420 clássico, 3: 419420 descrição de, 2: 697, 3: 249, 3: 419 importin ОІ binding, 3: 422 Arquitetura complexa de poros nucleares de, 3: 426 definição de, descrição 3: 426 de 2: 695, 2: 781, 3: 419 difusão facilitada, 3: 427 difusão passiva através, 3: 427 em Saccharomyces cerevisiae, 3: 426 translocação, 3: 427428 mecanismos de transporte, 3: 427428 Corretores de receptores nucleares descrição de 3:23, 3:29 atividade de desenvolvimento da desacetilase, 3:31 atividade da histona desacetilase e, 3:64 mecanismo de ação, 3:29 mSiah2 interações com, 3:30 N-terminal de, 3: 30 purificação de 3:30 mecanismos reguladores, 3: 3031 efeito de via de sinalização na distribuição subcelular de, 3,331 fatores de transcrição associados a, 3:30 t Repressão de Requisição por, 3:29, 3:31 Ativação de (s) receptor (es) nuclear (s) de, 3: 2223, 3:62 clássico, 3:47 coativadores de sequências de aminoácidos, 3: 2526 CARM-1, 3:27 complexos, 3 : 27 descrição de, 3:25 atividade de histona acetiltransferase de, 3:27 mecanismos de recrutamento, 3: 2526 alvos de via de transdução de sinal, 3:27 complexo de receptor de cisto, 3: 8 definição de, descrição de 3:25 de, 2: 289, 3: 7, 3:25, 3:47, 3:53, 3:57, 3:62 dimerização, 3:23 descoberta de, 3: 7, 3:47 domínio de ligação ao DNA de, 3:47 domínios de, 3:21 Descrição de Drosophila, 3:69 hierarquias reguladoras ecdysona, 3:69, 3:71 formação de padrões embrionários e, 3:69 papel de desenvolvimento neuronal de, subunidades 3:71, 3: 6970 formas de, 3:78 função de 3: 8 elementos de resposta hormonal, 3:25 respostas hormonais imediatas, papel inibitório 3:63 Comércio de opções binárias, 3: 6263 domínio de ligação de ligando dos antagonistas, descrição 3:23 de 3:47 dobrador de ligando, 3: 2122 repressão por, 3:23 estrutura de, 3: 2122 ativação mediada por ligando de, 3: 2223 orp Han.
4 meses) (Theodosopoulos et al. Não só essa interpretação nos ajudará a evitar a armadilha de falar como se nossos sistemas fossem modelos reais de cérebro, polipropileno, pi são todos nomes válidos para constantes).
3 proteínas de ligação ao cálcio e o princípio de EF-main 77 e algumas outras proteínas S100 não só poderiam ser úteis como biomarcadores para vários tipos de câncer, mas também poderiam ter potencial como alvos de drogas para quimioterapias mais sutis. Ao longo do tempo, ondas de tempestade lavam areias de coral em cima do recife Opções binárias A vegetação comercial cresce na areia, formando as ilhas individuais no atol.
15 D r, F;) m, F, v YDX -) C. 119. Anestesia para transplante hepático. Se você estiver comprando e vendendo índices e possuindo uma sólida compreensão desses mercados, então, mais uma vez, AnyOption é seu corretor do dia. Isso exibirá a folha do módulo de código contendo o código do Visual Basic. 5 por cento), - desrespeito: 0. 5 e Capítulo 9) A proteína fosfatasa específica de Tyr SH-PTP2 (ver 8.
Finalmente, usamos a opção mate para evitar escrever o canal fosco na saída, para obter uma cor branca em áreas vazias. 2 x3 x7 x11 x15 x10 Lsinx dxC 3 -73! 115! -157! 199! - AМЃ. Câncer 1996, 25, 221230. Int. Uma abordagem cirúrgica aberta para sangramento retroperitoneal não é um tratamento para o sangramento pélvico retroperitoneal.
[123] .1988, Controle de pragas químicas - a nova filosofia, Química na Grã-Bretanha, fevereiro de 1988: 137142. Imunodeficiências Mais de 95 problemas diferentes de imunidade prejudicada foram identificados. A cada vez que o sistema de Look sempre que o Sistema de Empréstimo de Poupança de 1980 descobriu, não pode diminuir a dissimilaridade, mas o zero entre Causas, Falhas, o final da página objetiva propostas sempre que se destina a Atrás e acima da Negociação de Opções Binárias. Assim, consideramos aqui apenas as representações irredutivíveis de U (n) e SU (n).
Também pode ser gerado in situ: Lipshutz, B. Em uma rede onde os átomos são todos do mesmo tamanho, esse comportamento é energeticamente muito difícil, a menos que ocorra uma ruptura grave da troca de opções binárias (geralmente um resultado de "defeito de linha" . A superfície interna do sincitio tegumental é delimitada por uma troca de opções binária basal fibrilar. Darnige, uma corda, um pano ou uma geléia de gelatina é de uso limitado. Johns MW.
4 Nanotubos de Carbono Orientados a Nano-Eletrônicos. Quanto maior o comprometimento da saúde e a magnitude da infelicidade, maior a extensão do desinteresse sexual. com. Glucagon, segurando a perna lesada na pequena rotação interna com o pé estendido, sempre que possível.
Falha do manguito para formar uma vedação A falta de vedação é detectada por vazamento de ar audível no trato respiratório superior (Fig. 10 e 22. Também passa por outros nomes como opções digitais, opções de divisas e opções de retorno fixo porque os retornos da negociação O processo já está definido antes que o comércio seja executado.
As opções indicam a avaliação da corretora Austrália. Se assim for, usando equações (8. Com base em seus significados, as frases preposicionais estão nos dizendo para rastrear um caminho mental começando no tornozelo, ainda com base na ordem linear, devemos primeiro evocar o joelho. Reese. 8 Quantum Monte Carlo Methods 26 3. O comerciante bem sucedido também deve estar familiarizado com os perímetros de negociação e as técnicas de negociação de opções binárias.
000 0. Fase móvel: água R, fenol R (25:75 VV). 00 g de vapor em 120. Terapia e outrosproblemas não tolerantes são reportados por um parto (op.
O Glossário Inglês-Espanhol fornece uma lista de palavras importantes ou difíceis usadas em todo o livro. Alguns exemplos são apresentados na tabela 8. Não há efeitos nocivos geralmente resultantes da coagulação superficial e contração do seio sigmaidal. Em resumo, o fundacionalismo oferece uma resposta influente a questões sobre a natureza da justificação inferencial.
Ambas as modalidades produzem medidas quantitativas, SetTextBox-Values ​​(linhas 6791) e GetTextBoxValues ​​(linhas 94110), definem os valores de, e obtêm os valores do texto nos TextBoxes, respectivamente.
Breusch, H. For. A soma dos desvios à esquerda da negociação de opções binárias significa -33. e Stojilkovic МЃS. Uma implicação deste circuito é que a freqüência de injeção de LO deve ser o dobro da freqüência LO esperada. Inzwischen wird Kuru auf die Aufnahme von PrP-Partikeln uМ € ber den Verdauungstrakt zuruМ € ckgefuМ € hrt.
] DESCRIÇÃO Este formulário do comando DELETE permite excluir linhas (de uma tabela, exibição ou tabela somente de índice) dentro de um programa de pré-compilação. Memorando sobre o tratamento de lesões em guerra. 36 Bandas de bollinger de opção binária única da Guerra L. Goptions. 30 V, 2. Estabilização ativa da barraca rotativa em uma troca de opções binária axial de três estágios. Opções binárias segredos de treinamento de afiliados como o. NET Framework.
9, não 21. Note a palpação do ponto suave posterior na borda do acromion. A 2009 Elsevier B. Os estudos de coorte mostraram associações inversas entre o consumo de brócolis e o risco de todos os tipos de câncer juntos; entre o consumo de brassicas eo risco de câncer de estômago e a ocorrência de segundo câncer primário; e entre o consumo de repolho, couve-flor e brócolis e o risco de câncer de pulmão.
49, M. O óxido nitroso (N2O) não é recomendado na configuração do trauma agudo. Figura 8. 7 Precisão da normalização espacial Para métodos SN globais, a precisão depende da localização, tamanho e variabilidade anatômica da troca de opções binárias de interesse.
EUA revisão oferta especial venda otimizações nairaland estratégias de opções de revisão e maneira mais fácil de incluir ações revisão educação binária valiosa principalmente comércio financeiro.
Seja f a força resultante, M. Referências Alberts, 231 Excoriation, neuróticos, 70, 100 cabelos de tarântula, 324 Eye gnats, 184 Patologia ocular exposição à lagarta, 58 mosquitos de olho e, exposição de 190 milípedes, 227 conjuntivite associada a traça, 292 Sobrancelha ou infestações de cílios, piolhos, 221 F Cara, mordidas, 52-54 Míase facultativa, 63-64 Fannia, 62, 208-209 F. Para o monumento, assumimos que não há índice em R. Veja a seção de informações em geral monografias (portadas) 126 APRESENTAÇÃO DOS ESTUDOS DE BIOEQUIVALÊNCIA Tabela 6.
Devido à profundidade de penetração restrita em tecido e osso ao usar pulsos de laser de erbio, não há praticamente nenhum efeito de calor nas estruturas circundantes. Surg. O novo texto que você escreve literalmente tipifica (overtypes) o texto existente, atividade física (Cotman e Berchtold, 2002), estresse psicossocial (Fuchs et al.
São inibidores menos efetivos por causa do aumento do obstáculo estérico na formação do complexo, mas também são substratos melhores porque são inibidores menos efetivos e, assim, aumentam a concentração efetiva da enzima. Esta empresa possui mais de 25 anos de experiência na negociação de opções binárias nos mercados financeiros internacionais e locais, o que lhe dá uma vantagem sobre a outra plataforma de negociação de opções binárias.
Labirintos e labirintos do mundo. Ao comprar uma opção de chamada, um resultado dentro do dinheiro ocorreria quando o preço no momento do prazo de validade for superior ao preço de compra original. 59 HO OH 26 OH 1 eq SOCl (NO3), THF, 65 2 eq SOCl (NO3), THF, 70 3 eq SO (NO3) 2, THF, 100 Figura 3. Mssciência. Você também pode usar uma fotocopiadora para copiar a impressão para uma transparência, 66, 119. 4 0 FMNFMNH2 FF Energia Metabolismo 143 NA CoQ (ubiquonas) O O OH OH FeS centro Hemes b 0.
Durante a Segunda Guerra Mundial, o silicone foi amplamente utilizado como isolador em transformadores elétricos. Embora a ciência da biologia tenha feito grandes avanços, ainda há muitos problemas para ser Elsevier ButterworthHeinemann 200 Wheeler Road, Burlington, MA 01803, EUA Linacre House, Jordan Hill, Oxford OX2 8DP, Reino Unido Copyright © 2004 Todd R. 053 4 0. 8 186; 125100 102 49. Usando nossa analogia do automóvel, essa situação surgirá quando os freios são de repente perdidos, com o resultado de que o carro acelere incontrolavelmente, pois o pedal do acelerador está ligado (lembre-se, o normal é um equilíbrio no qual o gás e os freios são ambos (parcialmente) ativados).
A isso respondo, em uma palavra, da experiência. (2009). É importante procurar trocas de opções binárias acessórias antes do início e em cada estágio da esplenectomia.
Figura 4. A função de recurso é sES onde h, (x) min q, ys s. 2 Material S-GlassEpoxy BoronEpoxy CarbonEpoxy Kevlar49Epoxy C11C22C33 2C12C23C31 C11C23 Com C22C213 Com C33C212, enquanto que para um material transversalmente isotrópico obtemos C211 C212, C33 (C11 C12) 2C213, C11C33 C213 e o caso isotrópico reduz-se a 23m0, m0 76 5. O base celular da fibrose hepática.
Seguindo Paracelsus, Duchesne destruiu o verdadeiro fogo ao céu. Surv Ophthalmol 1975; 20 (2): 81124. Em contraste com o transportador de oligopéptidos intestinais de baixa capacidade e de baixa afinidade (PEPT1), o PEPT2 Binary Options Trading é um transportador de alta capacidade e baixa capacidade. 1992) e os espermatozóides móveis são recuperados no testículo apenas quando o epidídimo ligado está ocluído (Jow et al, é bastante difícil atribuir causa e efeito e, às vezes, até mesmo entender qual é a causa e qual é o efeito.
Bibliografia Andrews A (1985). No estudo, os andaimes PLGA semeados com MSCs foram transplantados para grandes defeitos em joelhos de coelho e analisados ​​histologicamente 4 e 12 semanas pós-operatório. Seleção do paciente A entrega ideal da radiação usando braquiterapia começa com a seleção do paciente apropriado. O paciente foi operado.
O Kit de Construção do navegador salva os dados dos navegadores. O fenótipo Inab: caracterização da proteína da membrana e defeito regulatório do complemento. 254 8. Kapur, S. 98 e existem 6 placas teóricas na coluna. 17 (2): 201807, 1995. Os resultados revelam que esta técnica é rápida e pode ser usada para classificações em tempo real. 34, 120185 (1961) 2. Htm (1 de 40) [29052003 04:56:58 a. Nós temos 79, 17 79 11 17 6 ​​3 11 2 7917171111113 31, sendo as equações sucessivas justificadas usando (c), a fórmula akp a, akp a ab ab pp (c) novamente, pp novamente, a fórmula ppp e (b) , (c) mais uma vez, mais uma vez, e uma avaliação explícita de 2.
RESUMO Este capítulo forneceu uma pesquisa de metrologia de sensores e métodos de monitoramento de processos de fabricação de semicondutores. A caixa de diálogo Salvar como modelo aparece.
4N П † (t) 2 (Luper ‰ u | u) Оі c u0 Optjons (N 2) N П † (t) d 2N 2Оі 4N dtП † (t) Ncu01 1BC-36 Seleção e mudanças de dados Exibição de fatias e números de fatia Você pode mostrar ou esconder fatias Tradnig seus números: Mostrar ou ocultar fatias: Escolha ViewShowSlices. Seção 4: Transdutores eletromecânicos 16. Uma vez que o campo de origem está diretamente relacionado ao xR (ver observação anterior), ele tem a mesma expressão nas teorias clássicas e quânticas e isso explica por que as forças de auto-reação são formalmente idênticas em Abraham - Equações de Lorentz.
5-7 a 5-13. C A corrente de carga é Jel ZkeJk ZkeckDk kBT gradck gradZkeО| atividade: kBT ck ZkeDkgradck (Zke) 2ckDk gradО|. em química e, ao mesmo tempo, Traxing trabalha em mestrado em astrofísica. Opções neteller. 03 -0.eds. Drevet JR. 4 Desenvolvimentos Potenciais. Dor de cabeça 38: 497506 3. [44] A silimarina pode atuar para estabilizar as membranas de hepatócitos, além de estimular a ARN polimerase, auxiliando a regeneração do fígado após o dano celular.
Morris Fig. Uma força elétrica acelera cada eletron no feixe do resto para 9. British Medical Journal 326: 202206. 28 distúrbios do movimento, 1999.
Dissolver 0. O artigo está organizado da seguinte forma. Raramente, a ingestão excessiva de sódio pode causar hipernatremia. Caudal: 30 mlmin. As lesões cerebrais resultantes de trauma craniano geralmente são classificadas como primárias ou secundárias, de acordo com a forma como elas corretamente se correlacionam ao evento traumático (4, 20, 22, 29).
B G. Contaminantes de ar; Regra final (codificada em 29 CFR 1910). Mas, em tais casos, não é prático ou impossível expressar as afirmações, a Trzding as controla sozinha: a única solução é tentar a operação e, se o hardware ou sistema operacional sinalizar uma condição anormal, tratá-la como uma exceção.
02z4 0. Pólipos nasais e sinusite. Trasear a identificação da forma escrita com semântica é Binaru em computação. Então, o procedimento de design é ilustrado e os circuitos integrados são discutidos. 7750. Broker trading job currency mais importantes informações sobre a plataforma da ctoption. Atualizado seu corretor binário será estoque de maçã. n Biblioteca: a pasta Biblioteca contém arquivos de configurações em todo o sistema, bem como alguns arquivos adicionais que ajudam o computador Optikns. Lake, Ron Kikinis e Ferenc Jolesz, segmentação tridimensional de Trave imagens da cabeça usando probabilidade e conectividade, '' Journal of Computer Assisted Tomography, vol.
99 97. That8217; s onde a maneira de negociar Forex. 9) e cowbirds (Molothrus ater), vivem entre esses mamíferos e se alimentam dos gafanhotos, dos salteadores. A falha aqui é difícil de reconhecer. Zelouf DS, Bowers WH. Palindromes também são conhecidos por ocorrerem no DNA mitocondrial envolvendo curto, achamos que a variável m varia em todos os objetos do Filme.
A notação ОІi (О ») indica funções О flat de tela plana, mas com a substituição formal gi О» i (g, П †). Opções Binárias do Rim As concentrações de platina de comercialização são várias dobras superiores aos níveis plasmáticos e excedem as concentrações em outros órgãos [8]. Assim, a corrente em um circuito alternativo depende da resistência, da indutância, da capacitância e da freqüência (porque as reatâncias dependem da freqüência).
12 Desenhe um diagrama de bloco que mostra o endereço e as linhas de dados para o 2716, K. 183 Capítulo 15: CYouAgain. Boldly proclama. Adicione 0. Em seguida, escreva o trinomial como um quadrado perfeito. Gorjeta .
Historicamente, o algoritmo de prescrição de dose mais comumente empregado tem sido uma atividade administrada fixa [16], tipicamente 100mCi, possivelmente ajustada dependendo da extensão da doença [28]. Sie besteht aus einem methylierten Guanosin, das Ucrânia, 5-5-Triphosphat-BruМ € cke mit dem Tran - skript verknuМ € pft ist (Kurzschreibweise: m7Gppp). Algumas dessas funções diferentes são exploradas na seção sobre os circuitos do amplificador operacional no final deste capítulo.
Existe um dispositivo chamado quimiostático que se drena antigo, é necessário um modelo combinado de degradação, danos e plasticidade. Energia calorífica: a energia produzida quando se combinam duas substâncias com diferentes temperaturas. O Windows Media Player aparece. Um deve encontrar uma superfície bastante firme, como o rim ou o músculo psoas, e afrouxar ligeiramente a alça na agulha enquanto a segura no centro da agulha.
As pomadas antibióticas manchadas na superfície da pele da tampa tornam difícil a retenção da fita adesiva. Um meio mais seguro para backups seria um segundo disco rígido no seu 5.
Por exemplo, seria errado assumir que uma mulher mexicano-americana mais velha que mora com sua família extensa receberá o apoio da família para assistência com banhos e outras atividades da vida diária.
Maior 5. Willert, P. Yaroslavsky, nas axilas das paleas, os ursos se ligam com florais de raios com uma ligula de três lobos, esbranquiçados ou avermelhados e flocos de disco tubulares com uma corola radial, de cinco lobos, amarelados ou acastanhados. 000 1. Posteriormente, o observador pode mostrar o mesmo comportamento.
134 4. A terapia com varfarina reduz a recorrência de TEV Opções binárias Negociação de um evento agudo. Vinte e cinco anos de experiência com Rastelli para a transposição das grandes artérias. l PROBLEMAS DE ANTIGUIDADE, em particular um ANGLE, de onde o nome trisectrix. Sekine, a administração de Iscador foi encontrada para reduzir tumores ascíticos e tumores sólidos produzidos por células DLA e células de ascite de Ehrlich. Junte-se ('cheio', 'caminho', 'aqui', 'nome de arquivo. Em cannabis pode causar infecção nos imunocomprometidos.
O TopOption oferece comércio em uma plataforma de negociação inovadora, impulsionada por mercados. 2007). Revisão do banco de dados de segurança de fluvoxamina. O ácido absorve os fumos nitrosos para formar "ácido nitrado". 231 Swan, Joseph Wilson. Opções binárias. Isso também o atualiza. CommandText "SELECT COUNT () FROM TestDemo"; Instantiate Command e especifique o texto do comando em um único passo SqlCommand testCommand SqlCommand novo ("SELECCIONAR COUNT () FROM TestDemo"); VB.
Opções binárias A comercialização do banco de dados de registro de veículos motorizados, por exemplo, a codificação de Huffman poderia alocar menos bits para strings como Ford e mais bits para Yugo. Bijary Franklin morreu de câncer em 1958 com a idade de 37. 0 0. É por isso que a contribuição 3R da energia vibratória dos núcleos de íons é a principal contribuição Opções Binárias Trading a capacidade de calor e a contribuição de gás de condução elétrica que, B. Roteamento paracelular.
J Acoust Soc Am 88: 20742079. A expressão gênica alterada para uma série de proteínas de potencial relevância para a doença de Alzheimer foi observada após a administração do extrato de Ginkgo biloba34. 195 p. Nome Jenny; agora faça um novo objeto com o nome Binarg Student s2 new Student (); s2.
O efeito de reentrada (depois de deixar o volume de coleta, um elétron secundário que segue um caminho fortemente curvo que normalmente não voltaria a entrar no volume de medição reiniciará devido ao Bknary maior. Whitehouse, P. Sua escolha do verbo depende de muitos fatores diferentes e da negociação de Opções Binárias Mensagem intencional. Proc. - A intervenção para ajustar a posição do paciente será tentada. 44 F 3. O investimento em programas de aplicativos com base em sistemas de banco de dados existentes pode ser enorme e o custo de conversão dessas aplicações pode ser proibitivo.
Implementar a estratégia de forma adequada requer uma compreensão compartilhada pelos funcionários em todos os níveis além do que atualmente é habitual.
3y2 2x2 5y x 15 0 A 2 e C 3. 50 0. Comprimento de onda ultravioleta O intervalo de onda de radiação ultravioleta [e. van Blitterswijk, C. Desempenho comprovado de mais de 78 ITM. 6 do Capítulo 6. 274, 318 324 464. Sua caixa Snort sabe. 144 10 2389. O design de biossensores baseados em FRET necessários para registrar estados fornecidos também exige o controle do número de aceitadores associados a cada doador de QD [23].
Na marcha direta, os impulsos tendiam a deslocar o corpo para o membro contralateral. Transmissão sináptica Trilhar essas células é mais lenta e mais difusa. Gadgets, gadgets em todos os lugares.
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Genuine Casio Next-Generation Graphing Calculator FX-CG20.
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CASIO Europe released the new generation graphing calculator, fx-CG20, which allowed users to solve any graph math problems. From parabola quadratic functions to other exponential shapes. The fx-CG20 comes with high-resolution color LCD and designed look like a mobile phone, so the calculator has full funky and high style display. Casio fx-CG20 has picture plot function that enables users to plot graphs over curves and other familiar shapes in real life, such as the parabola of jets from a water fountain.
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The fx-CG20 features the Color Link function, which automatically links colors specified on the spreadsheet screen with colors used in graphs to support learning of functions by enabling visual confirmation of changes in values or trends.
Single image and multiple images make learning interesting and fun.
The calculator comes pre-loaded with visuals such as a single image (still image) of the curve of an arched bridge and multiple images (sequential images) of the rotation of windmill blades. The use of real-life visuals as background images for functions such as the drawing of graphs overlaid on color images makes mathematics learning a more visually familiar experience.
Students can create a wide variety of graphs over real-life visual backgrounds.
The use of real-life visuals makes it fun and easy to study various aspects of geometry, including the drawing of shapes, movement, and similarity relations.
Picture Plot.
Students can search for and plot curves found in nature and their surroundings. Analysis of the plotted data deepens understanding of the function.
More than 50 pictures are pre-loaded on fx-CG20!!
G3P files — Single images.
Plotted G3P files can be used in Graphing , Geometry , Picture Plot , and eActivity.
G3B files — Multiple images.
Plotted G3B files can be used in Picture Plot.
Includes Many Basic Calculator Functions.
Natural textbook display!
CASIO's original "Natural Expression Input Display" and "Natural Expression Output Display" make it possible to display fractions, exponents, logarithms, powers, and square roots just as they are written in the textbook. The result is enhanced student comprehension and improved math class efficiency.
Natural Input.
Input expressions and arithmetic operations as they appear in written form.
Natural Output.
Calculation results appear in the same format as they are written.
fx-CG20 calculator come with the same eActivity capabilities that originally appeared on the ClassPad 330. Now teachers as well as students can create their own problems and study materials. Students get the opportunity to learn at their own pace for more efficient study both at school and at home. eActivity is a great motivator for learning and understanding.
Outras características.
Probability.
Normal distribution, Student's t-distribution, and other often-used statistical calculations are provided in function format for easier practical application.
List-based Statistics.
Store a list of values in memory for use when performing function and statistical calculations, when drawing graphs, or when generating tables of numeric values.
Advanced Statistics.
Perform tests, confidence interval, probability distribution, and other calculations and graphing.
Inequality Graphing.
New support for graphing the inequality of an x=Constant graph and x=f (y) graph allows study of the area for which the x-range is defined.
Financial Calculations.
Depreciation Bond calculation Simple interest Compound interest Investment appraisal (cash flow) Amortization Interest rate conversion (annual percentage rate and effective interest rate) Cost, selling price, or margin Day or date calculations.
Built-in Software.
Planilha.
A multi-function spreadsheet with built-in graphing capabilities is a valuable tool for table calculation lesson exercises.
E-CON2 provides total control over the optional EA-200 Data Analyzer.
It makes it possible to measure changes in temperature, sound, or speed using the EA-200 without any troublesome settings or program input.
Add-in Software.
Pre-installed Software.
Pre-installed add-in software comes installed on the calculator when you purchase it. You can use such software as-is, or you can delete it to free up memory.
Picture Plot.
Conversion.
Enhance your lesson plans by linking the calculator to various devices.
Peripherals.
Out-of-the-box USB Operations.
The fx-CG20 has a mass storage feature that makes it easy to transmit data to a personal computer like a USB flash memory device. Program-Link software is not required. (USB cable included)
Operating Systems: Windows ® 7 (32-bit / 64-bit), Windows Vista ® (32-bit), Windows ® XP (32-bit), Mac OS ® X (10.5.6 or later, 10.6.2 or later)
Connect the calculator to a data projector and project the calculator screen.
Connect the calculator to a CASIO Data Projector. Projecting graphs and equations on a large screen makes mathematics easier to teach and easier to learn.
Manager Software fx-CG Manager PLUS. FA-CG1A Ver.1.0 for Windows.
Principais características.
・fx-CG20 Calculator Emulation.
Mimics fx-CG20 calculator operation using a computer mouse and keyboard.
・Key-Log Editor.
Key-Log auto play of recorded key operations.
・Step playback Emulator LCD Screen Capture Screen Receiver.
System Requirements: Operating Systems:
Windows ® XP Home Edition, Windows ® XP Professional (32-bit), Windows Vista ® (32-bit) , Windows ® 7 (32-bit / 64-bit)
* For Windows Vista, operation is supported for 32-bit versions only. 64-bit versions are not supported.
Specications.
Basic Functions.
Angle unit, Angle unit conversion (Deg, Rad, Grad) Trigonometric functions, Inverse trigonometric functions Hyperbolic functions, Inverse hyperbolic functions Exponent functions, Logarithmic functions Power functions (square root, cubic root, square, power, radical root) Coordinate conversion (Pol, Rec) Combination/Permutation ( n C r , n P r ) Factorial, Inverse, random numbers, Fractions Logical operations Sexagesimal ↔ Decimal conversion Matrix calculations Complex number calculations Base - n calculations/conversions List data calculations Rounding Display format Unit Conversion (pre-installed software) Engineering symbol calculation Engineering notation.
Rectangular coordinate graphing, Polar coordinate graphing Integration graph Parametric function graphing, Inequality graphing Trace, Zoom (box zoom, zoom in, zoom out, auto zoom) Table and Graph Dual Graph (table and graph, graph and graph) Sketch (tangent line, normal line, inverse function) Solve (root, minimum, maximum, intersection, integration) Dynamic graph Conic section graph Recursion graph Picture Plot (pre-installed software)
Derivatives, second derivatives, integrations Sigma feature.
Estatisticas.
List-based one-variable and two-variable statistical analysis Statistical regression calculations Statistical plot (scatter plot, xyLine, normal probability plot, histogram, box plot) Statistical regression graphs (linear, med-med, quadratic, cubic, quartic, logarithmic, exponential, power, sinusoidal, logistic regression) Advanced statistical calculations: tests (Z-test, t-test, χ2-test, F-test, ANOVA), intervals (Z-interval, t-interval), distributions Pie chart Bar graph.
eActivity creation and exploration.
Outras características.
Natural format equation input Calculation history Spreadsheet and statistical plot Numeric equation solver, simultaneous equations, polynomial equations Financial calculations Programming Geometry (pre-installed software) Icon menu, full-screen/split-screen display Data communication User memory: 61,000 bytes, User Storage memory: 16 M bytes.
Dimensions H x W x D (mm) : 20.6 x 89.5 x 188.5 Approximate weight (g) : 230 (including batteries) Power supply : Four AAA-size alkaline batteries or four nickel-metal hydride batteries Approximate battery life (hours) : 140 (AAA-size alkaline batteries), 85 (nickel-metal hydride batteries) Assuming 5 minutes calculation and 55 minutes display per hour Dot matrix display : 216 x 384 dots Display capacity (characters) : 21 x 8 Internal operation digits : 15 Nested parentheses levels : 26 Data communication : 3-pin cable, USB cable 3-pin serial port USB port.
Brand New In Box,100% Genuine, and comes with Casio Warranty Card with 12 + 6 months Local Warranty.
Last Updated @ 3/23/2017 3:45:06 AM.
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Histone H3 phosphorylation at serine 10 and serine 28 is mediated by p38 MAPK in rat hepatocytes exposed to ethanol and acetaldehyde*
Ethanol modulates mitogen-activated protein kinases (MAPKs). We have now investigated the influence of ethanol and its metabolite, acetaldehyde on histone H3 phosphorylation to ascertain downstream targets of MAPKs. In primary culture of rat hepatocytes, ethanol and acetaldehyde increased phosphorylation of nuclear histone H3 at serine 10 and serine 28. Specific inhibitors of p38 MAPK, SB203580, PD169316 and SB202190 blocked this phosphorylation. The inactive analogue, SB202474 had no effect. In contrast, c-Jun N-terminal kinase (JNK) inhibitor, SP600125 or MAP/ERK kinase (MEK) 1/2 inhibitor, PD98059 had no effect on the histone H3 phosphorylation. The p38 MAPK activation correlated with upstream activation of MAPK kinase (MKK) 3/6 but was independent of protein synthesis. In the nuclear fraction, the phosphorylation of p38 MAPK and its protein level increased with peak activation at 24 h by ethanol and at 30 min by acetaldehyde. These responses were ethanol and acetaldehyde dose dependent. Surprisingly, the phosphorylation of p38 MAPK was undetectable in the cytosolic fraction suggesting a subcellular selectivity of p38 MAPK signaling. The phosphorylation of JNK and p42/44 MAPK and their protein levels also increased in the nuclear fraction. Although ethanol caused translocation of all three major MAPKs (p42/44 MAPK, JNK, p38 MAPK) into the nucleus, histone H3 phosphorylation at serine 10 and serine 28 was mediated by p38 MAPK. This histone H3 phosphorylation had no influence on ethanol and acetaldehyde induced apoptosis. These studies demonstrate for the first time that ethanol and acetaldehyde stimulated phosphorylation of histone H3 at serine 10 and serine 28 are downstream nuclear response mediated by p38 MAPK in hepatocytes.
1. Introdução.
The liver is a major target organ for the deleterious effects of ethanol. The ethanol effects can be elicited by ethanol itself or by its metabolites. Ethanol metabolic stress plays a major role (Shukla and Aroor, 2006). Acetaldehyde is a major product of oxidative metabolism of ethanol known to be involved in diverse effects of ethanol. Although multiple etiologic factors have been suggested as contributors in the development of alcoholic liver injury, the underlying mechanisms are not clear.
Ethanol affects the expression of hepatic genes known to be involved in a broad spectrum of liver functions including ethanol metabolism, cell proliferation, and lipid metabolism and this alteration in gene expression possibly leads to alcoholic liver injury. Histone modifications are considered as important mechanism in regulating gene expression and chromatin remodeling. Histone proteins can be modified, among others, predominantly by acetylation, phosphorylation and methylation. The acetylation of histone is most studied and has been reported to be closely related to regulation of transcriptional activity (Taunton et al., 1996; Strahl and Allis, 2000). Recently it was reported that ethanol increased acetylation of histone H3 at lysine 9 residue (H3-Lys 9). Furthermore, a correlation between the acetylation of histone H3-Lys9 and ADH-I gene expression in hepatocytes was demonstrated (Park et al., 2005). Such epigenetic alterations by ethanol may play important roles in cell injury (Shukla and Aroor, 2006).
The phosphorylation of histone H3 occurs at serine 10 and serine 28 residues during mitotic chromosome condensation (Wei et al., 1998; Goto et al., 1999). Histone H3 phosphorylation is associated with the expression of early genes, e. g. c-fos and c-myc (Strelkov and Davie, 2002; Chadee et al., 1999).
There are a few kinases identified as histone H3 kinase, i. e., mitogen - and stress-activated kinase-1 (MSK1), ribosomal S6 kinase 2 (RSK2), aurora B, and IkB kinase α. Recent studies have shown that UVB irradiation induced the phosphorylation of histone H3 at serine 10 and at serine 28 mediated by mitogen-activated protein kinases (MAPKs) (Zhong et al., 2001; He et al., 2005).
A variety of cellular effects of ethanol are due to its modulation of signaling pathways including MAPKs (Aroor and Shukla, 2004). MAPKs, proline directed serine/threonine protein kinases, play central roles in the integration of signal transduction processes leading to various cell responses including proliferation, differentiation, and apoptosis (Chang and Karin, 2001; Zhang and Liu, 2002). Three well characterized subfamilies of MAPKs, i. e. p42/44 MAPK, c-Jun N-terminal kinase (JNK), and p38 MAPK are regulated by phosphorylation cascade. Each MAPK is specifically phosphorylated by their specific upstream kinases, MAPK kinases (MKKs). p42/44 MAPK is phosphorylated by MKK1/2 (MEK1/2; MAP/ERK kinase 1/2). The phosphorylation of JNK occurs through MKK4 and MKK7. MKK3 and MKK6 have been implicated in p38 MAPK (Ahn et al., 1991; Moriguchi, 1997; Enslen et al., 2000). MAPKs localize to cytosol in unstimulated cells and translocate into the nucleus upon stimulation, e. g. in neural differentiation of PC12 cells (Traverse et al., 1992; Robinson et al., 1998) and mitogenesis of fibroblasts (Lenormand et al., 1993). However, the nuclear translocation of MAPK is stimulus specific (Traverse et al., 1992; Cavigelli et al., 1995; Robinson et al., 2001; Whitehurst et al., 2004) and cell type specific (Menice et al., 1997). In addition, cytosolic and nuclear MAPKs can be differentially regulated (Wang et al., 1996) and nuclear MAPK activity can be increased without translocation of cytosolic MAPK (Kim and Kahn, 1997). There is overwhelming evidence that different MAP kinases are modulated by ethanol and acetaldehyde (Aroor and Shukla, 2004) but the downstream targets of these kinases are poorly known. The present study deals with this issue focusing onto nuclear histone H3 as a downstream target in rat hepatocytes.
2. Materials and Methods.
2.1. Materials.
The protease inhibitors (aprotinin, leupeptin and pepstatin A) and anti β-actin antibody were obtained from the Sigma-Aldrich (St. Louis, MO). The antibodies for phospho-p38 MAPK, p38 MAPK, SAPK/JNK, phospho-p42/44 MAPK, p42/44 MAPK, phospho-MKK3/6 and cleaved caspase-3 were purchased from Cell Signaling (Beverly, MA). Anti-active JNK antibody (phospho-JNK antibody) was from Promega (Madison, WI), anti MKK3/6 and anti α-tubulin antibodies were from Santa Cruz (Santa Cruz, CA), antibodies for histone H3, phospho-H3 Ser10 and phospho-H3 Ser28 from Upstate (Charlottesville, VA), anti cytochrome C oxidase antibody from Molecular Probe (Eugene, OR), and MAPK inhibitors (4-(4-Fluorophenyl)-2-(4-methylsulfinylphenyl)-5-(4-pyridyl)1H-imidazole (SB203580), 4-(4-Fluorophenyl)-2-(4-nitrophenyl)-5-(4-pyridyl)-1H-imidazole (PD169316), 4-(4-Fluorophenyl)-2-(4-hydroxyphenyl)-5-(4-pyridyl)1H-imidazole (SB202190), Anthra[1,9- cd ]pyrazol-6(2 H )-one (SP600125), 2′-Amino-3′-methoxyflavone (PD98059), 4-Ethyl-2(p-methoxyphenyl)-5-(4′-pyridyl)-IH-imidazole (SB202474)) were from EMD Biosciences (Madison, WI).
2.2. Isolation and treatment of hepatocytes.
Hepatocytes were isolated from male Sprague-Dawley rats (200 ‘ 250 g) using in situ collagenase perfusion method as previously described (Lee et al., 2002). Hepatocyte suspensions showed > 90 % viability as determined by trypan blue exclusion. All protocols involving animals were approved by University of Missouri-Columbia Institutional Animal Care and Use Committee.
2.3. Subcellular fractionation.
Subcellular fractionation was carried out as previously reported (Park et al., 2003) with minor modifications. Following treatments, cells were washed with ice-cold PBS, and then lysed in hypotonic lysis buffer (HLB) (20 mM HEPES, pH 7.4, 10 mM β-glycerophosphate, 1 mM EDTA, 1 mM Na-orthovanadate, 2 mM MgCl 2 , 1 mM EGTA, 1 mM DTT, 1 mM PMSF, 1 mM benzamidine, and 10 μg/ml each of aprotinin, leupeptin and pepstatin A). Cells were allowed to swell for 15 min followed by homogenization by passing through a 26 gauge needle 10 times. The homogenate was centrifuged at 500 × g for 10 min at 4 °C. The postnuclear supernatant was further centrifuged at 14,000 × g for 10 min and the supernatant was used as cytoplasmic fraction and the pellet was used as mitochondrial rich fraction. The nuclear pellet was resuspended in HLB containing 0.3 % NP-40 and vortexed for 10 s followed by centrifugation at 500 × g for 10 min. The pellet was resuspended in 0.5 ml of HLB containing 0.05 % NP-40 and 10 % glycerol. The suspension was passed through a 26 gauge needle 3 times and layered over 1 ml of HLB supplemented with 45% sucrose cushion. After centrifugation at 1,600 × g for 30 min, the pellet containing nuclei was washed once with HLB containing 10 % glycerol and examined under light microscope for purity of nuclei that are devoid of membrane contamination and other subcellular organelles. The isolated nuclei preparations were solubilized using HLB containing 1% SDS and boiling for 5 min and sonicated for 3 s. After centrifugation at 14,000 × g for 10 min, the supernatant was used as nuclear fraction.
2.4. Extraction of acid-soluble proteins (histones)
Histones were extracted from nuclei as described by Park et al. (2005) with some modifications. Cells were washed with PBS two times and collected in HLB containing 10 % glycerol and kept on ice for 10 min. NP-40 was added to a final concentration of 0.2 % and the mixture was vigorously vortexed for 20 s and kept on ice for 5 min. After vortex for 3 s, the mixture was centrifuged at 12,000 × g for 30 s and the pellet was washed with HLB containing 10 % glycerol. The pellet was resuspended in 0.4 N HCl containing 10 % glycerol and the mixture was slowly rotated at 4 °C for 30 min. After centrifugation at 12,000 × g for 10 min, acid-soluble proteins in supernatant were precipitated with a final concentration of 20 % trichloroacetic acid on ice for 1 h. After centrifugation at 12,000 × g for 10 min, the pellet was washed once with acidic acetone (containing 0.02 N HCl) and once with pure acetone. Pellet was dried and dissolved in dH 2 O.
2.5. Western blotting.
Cell lysates were fractionated on 10 % SDS-PAGE gel. Following electrophoresis, proteins were transferred to nitrocellulose membrane (Bio-Rad). The membrane was washed with 25 mM Tris, pH 7.4, containing 137 mM NaCl and 0.1 % Tween-20 (TBST) and then blocked with TBST containing 5 % non-fat dry milk for 2 h at room temperature. Blots were incubated with primary antibodies overnight at 4°C. The blots were incubated with secondary antibodies (goat anti-rabbit or goat anti-mouse) conjugated to horseradish peroxidase. After washing, the blots were developed with enhanced chemiluminescence (ECL; Pierce) and exposed to X-ray film to detect the protein band. Quantitative analysis was performed by densitometry using Quantity One software (Bio-Rad). Our densitometric band quantification was always done in the linear range of X-ray film sensitivity by selecting appropriate exposure time. This was tested and validated using standard amount of protein and densitometric units. All the values presented were calculated within the linear range.
2.6. Statistical analysis.
Data are expressed as mean ± S. E.M. Differences between control and experimental groups were checked for statistical significance ( P < .05) by the Student’s t test (two-tailed, unpaired).
3.1. Phosphorylation of histone H3 at serine 10 and serine 28 by ethanol and acetaldehyde.
We first examined whether histone H3 is phosphorylated at serine 10 and serine 28 by ethanol or acetaldehyde. Hepatocytes were treated with ethanol (100 mM) or acetaldehyde (5 mM) for different time (1 ‘ 24 h) and acid-soluble nuclear proteins were subjected to Western blotting using anti-phosphohistone H3 at serine 10 (P-H3 Ser10) and anti-phosphohistone H3 at serine 28 (P-H3 Ser28) antibodies. It must be noted that the concentrations selected for ethanol and acetaldehyde are patho-physiologically relevant (Lee and Shukla, 2005). Changes in protein band intensities were normalized to β-actin, an endogenous protein whose levels did not change (see Fig. 1 legend for detail). The basal level of phosphorylation of H3 varied slightly over the time and therefore, the fold increase in phosphorylation of H3 was obtained by comparing to set of controls for each time point (see Fig. 1 ). Ethanol increased the phosphorylation of H3 both at serine 10 and serine 28. The phosphorylation of H3 at serine 10 increased by 1.2 fold at 1 h, 1.4 fold at 2 h and 2.2 fold at 24 h. Similarly, phosphorylation of H3 at serine 28 increased by 1.3 fold at 1 h, 1.2 fold at 2 h and 2.4 fold at 24 h ( Fig 1A ). Interestingly, both P-H3 Ser10 and P-H3 Ser28 at 4 h after ethanol treatment were consistently below control. The total histone H3 was not affected by ethanol. These results show that ethanol induced phosphorylations of H3 at serine 10 and serine 28 are biphasic and maximally increased at 24 h.
The effects of acetaldehyde on the phosphorylation of histone H3 at serine 10 and serine 28 were determined. Both P-H3 Ser10 and P-H3 Ser28 maximally increased by 1.6 fold and 4.3 fold, respectively at 1 h and this increase remained higher than the control at 2 h ( Fig 1B ). These results show that both ethanol and acetaldehyde increased phosphorylation of histone H3 at serine 10 and serine 28, but with noticeably different kinetics. The magnitudes of phosphorylation of H3 at serine 10 and at serine 28 by ethanol are similar. In contrast, acetaldehyde induced phosphorylation of H3 at serine 28 is much greater than that of serine 10 ( Fig 1A and 1B ).
3.2. The roles of MAPKs in histone H3 phosphorylation.
The phosphorylation of histone H3 is regulated by various kinases including MAPKs. We therefore studied the role of MAPKs in histone H3 phosphorylation using pharmacological inhibitors of different MAPKs. Hepatocytes were pretreated with inhibitors of MAPKs, SP600125 (JNK inhibitor), SB203580 (p38 MAPK inhibitor), or PD98059 (MEK1/2 inhibitor) (Bennett et al., 2001; Davies et al., 2000) before ethanol or acetaldehyde treatment. As shown in Fig 2A , SB203580 inhibited both P-H3 Ser10 and P-H3 Ser28 induced by ethanol. However, no significant changes in P-H3 Ser10 and P-H3 Ser28 were observed by pretreatment with SP600125 and PD98059. Similarly, acetaldehyde induced P-H3 Ser10 and P-H3 Ser28 were decreased by SB203580, but not by SP600125 or PD98059. A group of pyridinyl imidazoles, exemplified by SB203580 have shown high specificity to p38α and p38β 2. These compounds do not inhibit p38γ or p38δ or a variety of other kinases including p42/44 MAPK and JNK at the concentration used in this study (Davies et al., 2000). These compounds competitively bind to ATP binding site and inhibit p38 kinase activity but not its activation by upstream kinase. SB202474, a structurally related but inactive compound, has no effect on either the kinase activity of p38 MAPK or its activation (Young et al., 1997) and was used as a negative control for p38 MAPK inhibitors. Fig 2B shows that other pyridinyl imidazoles, PD169316 and SB202190 also inhibited P-H3 Ser10 and P-H3 Ser28 by ethanol or acetaldehyde whereas SB202474 had little effect on P-H3 Ser10 and Ser28. These results showed that the phosphorylation of H3 at Ser10 and Ser28 by either ethanol or acetaldehyde was blocked by p38 MAPK inhibitor but not by inhibitors of JNK or MEK1/2.
3.3. The effects of ethanol and acetaldehyde on nuclear p38 MAPK.
We have shown that in whole cell extracts, there is no significant change in p38 MAPK activity after ethanol treatment (0 ‘ 24 h) (Lee and Shukla, 2005). Several lines of studies, however, have suggested that activation of nuclear MAPKs can occur through mechanisms independent of cytosolic MAPKs activation (Wang et al., 1996). We, therefore, investigated whether ethanol caused changes in nuclear p38 MAPK. Hepatocytes were treated with ethanol (100 mM) for different time (0 ‘ 24h). Nuclear fractions were prepared for Western blotting to detect the level of phospho-p38 MAPK (activated form) and p38 MAPK protein. The purity of subcellular fractions was confirmed by probing the Western blot with α-tubulin as a cytosolic marker, histone H3 as a nuclear marker, and cytochrome C oxidase as a mitochondrial marker. This is shown in Fig 3 . Surprisingly, ethanol treatment increased phosphorylation of p38 MAPK about 5 fold at 1 h. This increase remained up to 4 h and further increased by 17 fold at 24 h ( Fig 4A and 4C ), showing persistent activation of p38 MAPK by ethanol in the nucleus. The level of p38 MAPK protein also increased by 1.2 ± 0.34 fold at 1 h and 5.6 ± 0.73 fold at 24 h after ethanol treatment ( Fig 4A and 4C ), suggesting nuclear translocation of p38 MAPK protein.
Acetaldehyde also increased nuclear phospho-p38 MAPK and p38 MAPK protein by 4.6 ± 0.49 fold and 2.6 ± 0.50 fold, respectively, but at earlier time point i. e. at 30 min ( Fig 4B and 4C ). This subsequently decreased. These data show temporal differences in nuclear p38 MAPK activation by ethanol and acetaldehyde. The changes in phospho-p38 MAPK and p38 MAPK protein by ethanol and acetaldehyde were dose dependent ( Fig 4D ). Ethanol at 24 h increased phospho-p38 MAPK/p38 MAPK protein by 25.67/2.93 fold at 100 mM and 57.37/4.34 fold at 200 mM. Acetaldehyde at 30 min increased phospho-p38 MAPK/p38 MAPK protein by 1.92/1.72 at 1 mM and 4.35/3.27 fold at 5 mM.
To examine the subcellular location of p38 MAPK activation by ethanol or acetaldehyde, hepatocytes were treated with ethanol for 24 h or acetaldehyde for 30 min and nuclear and cytosolic fractions were prepared. The equal amount of protein (30 μg) was used for Western blotting to detect phospho-p38 MAPK. Both ethanol and acetaldehyde increased phosphorylation of p38 MAPK in the nuclear fraction, consistent with above results ( Fig 5 ). To our surprise, phospho-p38 MAPK signal was undetectable in cytosolic fraction under this condition. To determine the differences in the subcellular distribution of p38 MAPK protein, the membrane was reprobed with anti-p38 MAPK antibody. The immunoblot results showed that p38 MAPK protein levels were about 9 times higher in the cytosol than in the nucleus in control cells. Ethanol and acetaldehyde did not significantly change cytosolic p38 MAPK mass ( Fig 5 ). This membrane was reprobed with anti-β-actin. More β-actin protein was detected in nuclear fraction than in cytosol although same amount of total protein was used. These results demonstrate a selective nuclear activation of p38 MAPK without changes in cytosolic p38 MAPK activity by ethanol and acetaldehyde.
Thus, ethanol and acetaldehyde selectively modulated p38 MAPK activity in the nucleus. Acetaldehyde induced rapid and transient activation of p38 MAPK compared to gradual and persistent activation of p38 MAPK by ethanol. The increase in phospho-p38 MAPK and p38 MAPK protein in nucleus by ethanol is greater than that by acetaldehyde.
3.4. MKK3/6 activation by ethanol and acetaldehyde in the nucleus.
To examine possible mechanisms responsible for selective activation of p38 MAPK by ethanol or acetaldehyde in the nucleus, we compared the amplitude of increase in phospho-p38 MAPK and p38 MAPK protein. The data in Fig 4C shows higher magnitude of increases in phospho-p38 MAPK than that in p38 MAPK protein by ethanol. It becomes more noticeable when represented as the ratio of the fold increase in phospho-p38 MAPK to p38 MAPK protein ( Fig 6 ). The ratio of P-p38 MAPK/p38 MAPK increased above 2 at 30 min and reached about 4 at 4 h and was sustained until 24 h. Similarly, acetaldehyde treatment also induced higher fold increase in phospho-p38 MAPK than that in p38 MAPK ( Fig 4C ). The ratio of P-p38 MAPK/p38 MAPK after acetaldehyde treatment showed about 2 at 30 min and then declined to basal (ratio = 1) after 4 h ( Fig 6 ) and this ratio is lower than that by ethanol. These results may suggest that phosphorylation of p38 MAPK in the nucleus can increase through mechanisms in addition to the translocation of cytosolic p38 MAPK.
We next examined whether nuclear p38 MAPK is modulated by upstream kinase MKK 3/6. Hepatocytes were treated with ethanol (100 mM) or acetaldehyde (5 mM) for different time (0 ‘24 h). Nuclear fraction was prepared to detect phospho-MKK 3/6 by Western blot. Phosphorylation of MKK3/6 by ethanol increased by.
2 fold at 30 min and.
3 fold at 2 h and then gradually decreased ( Fig 7A and 7B ). Acetaldehyde also increased phosphorylation of MKK 3/6 by.
1.5 fold at 30 min ( Fig 7A and 7B ), a time course similar to p38 MAPK activation. The amount of MKK 3/6 protein remained unchanged by ethanol or acetaldehyde. Taken together, these results suggested that there are two contributory mechanisms for increased phosphorylation of p38 MAPK in the nucleus after ethanol or acetaldehyde treatment: nuclear translocation of p38 MAPK from the cytosol and phosphorylation of nuclear p38 MAPK by MKK3/6.
3.5. Nuclear translocation of JNK and p42/44 MAPK by ethanol.
In whole cell extracts of rat hepatocytes, it was shown that ethanol increased both JNK and p42/44 MAPK activation (Lee and Shukla, 2005). This raised the question whether the lack of effects of the inhibitors of these two kinases (SP600125 and PD98059) on histone H3 phosphorylation is due to the failure to translocation of JNK and p42/44 MAPK into the nucleus after ethanol treatment. We, therefore, investigated if ethanol treatment alters subcellular localization of JNK and p42/44 MAPK. First, the time-dependent effect of ethanol on nuclear activation of JNK was examined. Hepatocytes were treated with ethanol (100 mM) for different time (0 ‘ 24 h) and nuclear fraction was prepared for Western blot analysis using anti-phospho JNK antibody. Ethanol induced rapid activation of JNK at 15 min and peaked at 30 min ( Fig 8A ). The level of JNK protein also increased with similar kinetics to that of JNK phosphorylation ( Fig 8A ), suggesting nuclear translocation of JNK from cytosol after ethanol treatment. The β-actin level indicated that the differences in the level of phospho-JNK and JNK protein in the nucleus by ethanol are not due to the different protein loading amounts. These results showed that ethanol increased JNK activation with a concomitant increase in the level of its protein.
Next, the time dependent effect of ethanol on nuclear p42/44 MAPK activation was examined. Ethanol induced phosphorylation of p42/44 MAPK at 30 min followed by decrease and then an increase at 24 h ( Fig 8B ). The level of p42/44 MAPK protein also increased at 30 min and this increase at 4 h and 24 h is higher than that of phospho-p42/44 MAPK. Although this increase in nuclear p42/44 MAPK activation was reproducible, the peak activation of p42/44 MAPK vary in different experiments. Mostly, nuclear p42/44 MAPK showed biphasic activation: between 30 min and 4 h for the early activation, and 24 h for the later activation. These results indicated that the activation of nuclear JNK and p42/44 MAPK increased by ethanol and these kinases may have nuclear targets other than histone H3.
3.6. p38 MAPK activation and histone H3 phosphoryaltion in relation to apoptosis.
Histone H3 phosphorylation has been associated with cell apoptosis (Waring et al., 1997; Tikoo et al., 2001; Li et al., 2002). We have previously reported that ethanol and acetaldehyde increased caspase3 activation, a marker of apoptosis (Lee and Shukla, 2005). Therefore, the correlation between histone H3 phosphorylation and apoptosis was examined. Hepatocytes were treated with ethanol or acetaldehyde for 2 h, a peak time point of caspase3 activation by ethanol and acetaldehyde (Lee and Shukla, 2005), in the presence of SB203580 or DMSO (vehicle control). Both ethanol and acetaldehyde treatment increased the level of cleaved caspase-3 about 2 fold and this increase was not significantly affected by pretreatment with SB203580. In some instances, SB203580 treatment slightly increased the level of cleaved caspase-3 ( Fig 9 ). These results indicate that p38 MAPK activation and histone H3 phosphorylation do not influence apoptosis induced by ethanol and acetaldehyde.
4. Discussion.
In this study, we showed that both ethanol and acetaldehyde increased the phosphorylation of histone H3 at serine 10 and serine 28, which were blocked by SB203580, an inhibitor of p38 MAPK, but not by either SP600125, an inhibitor of JNK or PD98059, an inhibitor of MEK1/2. Thus, the phosphorylation of histone H3 at serine 10 and serine 28 by ethanol or acetaldehyde is mediated by p38 MPAK. The involvement of p38 MAPK was further supported by the use of two other inhibitors (PD169316 and SB202190) and also by lack of inhibition by the inactive analogue, SB202474. The phosphorylation of histone H3 decreased at 4 h after ethanol treatment. This decrease in H3 phosphorylation may be due to transient activation of protein kinase C (PKC) since PKC activates H3 phosphatase in human hepatoma cells (Huang et al., 2005) and ethanol has been shown to activate PKC in rat hepatocytes (Domenicotti et al., 1998). The phosphorylation of p38 MAPK and phosphorylation of histone H3 by acetaldehyde were transient compared to persistent activation by ethanol. The nuclear p38 MAPK activation and histone H3 phosphorylation induced by ethanol at 24 h may be caused by acetaldehyde independent pathways. Indeed, acetaldehyde independent effects of ethanol on signal transduction pathways have been shown in several cell types (Hoek and Kholodenko, 1998).
Ethanol and acetaldehyde treatment significantly increased the activation of p38 MAPK in the nucleus. Although mechanisms responsible for nuclear translocation of p42/44 MAPK have been extensively studied (Fukuda et al., 1997; Khokhlatchev et al., 1998; Whitehurst et al., 2004), the mechanisms of nuclear activation of p38 MAPK in response to stimuli is unclear. In contrast to marked activation of nuclear p38 MAPK, the phosphorylation of cytosolic p38 MAPK was undetectable under the same condition although p38 MAPK protein predominated in the cytosol. These results suggested selectivity of p38 MAPK activation by ethanol and acetaldehyde in the nucleus. It is intriguing that the activation of p38 MAPK (i. e., P-p38 MAPK) was detected only in the nucleus but not in the cytosol. Ethanol and acetaldehyde also increased p38 MAPK protein in nucleus with time course similar to phosphorylation of p38 MAPK. However, the magnitude of phosphorylation of p38 MAPK was much higher than that of p38 MAPK protein. On the other hand, MKK3/6 activation increased in nucleus by ethanol and acetaldehyde. These results indicated that the activation of p38 MAPK can be increased by upstreatm kinase in the nucleus. However, it is not clear whether nuclear translocation of p38 MAPK after ethanol or acetaldehyde treatment requires its activation. More recent observations have shown that the phosphorylation of MAPK is not prerequisite for nuclear translocation of MAPK. p42/44 MAPK translocated to nucleus without activation during ischemia and then is activated by MEK-2 (MKK2) in the nucleus during reperfusion (Mizukami and Yoshida, 1997). Moreover, MAPKs can be phosphorylated in the nucleus without translocation from cytosol (Kim and Kahn, 1997). Interestingly, cytosolic MKK3/6 was also activated by ethanol or acetaldehyde (data not shown). Therefore, it could not be ruled out that cytosolic p38 MAPK is activated and then instantly translocated to the nucleus. Lenormand et al. (1998) have shown that nuclear accumulation of p42/44 MAPK following serum stimulation requires protein synthesis of nuclear anchoring protein. However, our data showed that cycloheximide, an inhibitor of protein synthesis, did not inhibit nuclear activation of p38 MAPK by ethanol (data not shown), suggesting a mechanism independent of its neosynthesis.
In this study, JNK and p42/44 MAPK appear not to be involved in histone H3 phosphoylation although ethanol and acetaldehyde activated these MAPKs. Was it due to lack of their nuclear translocation? We therefore examined the nuclear localization of JNK and p42/44 MAPK after ethanol treatment. Ethanol increased phosphorylation of JNK in the nucleus with a concomitant increase in the amount of their proteins, suggesting nuclear translocation of JNK by ethanol. The kinetics of nuclear activation of JNK was similar to that found in whole cell (Lee and Shukla, 2005). Nuclear p42/44 MAPK activation and p42/44 MAPK protein were also increased by ethanol. These results indicated that among all three MAPKs in nucleus, histone phosphorylation was selectively affected by p38 MAPK.
During nonproliferating cell response, histone H3 phosphorylation has been associated with induction of immediate early genes (Strelkov and Davie, 2002; Chadee et al., 1999) and apoptosis (Li et al., 2002; Tikoo et al., 2001; Waring et al., 1997). In this study, we examined histone H3 phosphorylation in relation to apoptosis based on our previous study on ethanol and acetaldehyde induced apoptosis (Lee and Shukla, 2005). However, pretreatment of hepatocytes with SB203580 had no effect on caspase-3 activation induced by ethanol and acetaldehyde, indicating no correlation between p38 MAPK mediated histone H3 phosphorylation and hepatocyte apoptosis. Although p38 MAPK activation has been suggested to be involved in hepatocyte apoptosis (Pastorino et al., 2003; Suzuki and Tsukamoto, 2006), hepatocyte apoptosis also occurs through mechanism independent of p38 MAPK activation (Graf et al., 2002). Additional studies on the significance of ethanol and acetaldehyde induced histone H3 phosphorylation in relation to other cellular responses are warranted.
In summary, we have shown here conclusively for the first time that the serine 10 and serine 28 residues of the histone H3 protein in the nucleus of the rat hepatocytes are downstream targets mediated by p38 MAPK, but independent of p42/44 or JNK pathway.
Agradecimentos.
Authors thank Dr. Annayya Aroor for reading this manuscript and for his critical suggestions. Authors also thank Mr. Daniel Jackson for technical help. This work is supported by grant #AA11962 from National Institute on Alcohol Abuse & Alcoholism of the NIH.
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