DESENVOLVIMENTO EMBRIONÁRIO E DIFERENCIAÇÃO SEXUAL -/- E. I. C. da Silva Departamento de Agropecuária – IFPE Campus Belo Jardim Departamento de Zootecnia – UFRPE sede -/- 1.1 INTRODUÇÃO O sexo foi definido como a soma das diferenças morfológicas, fisiológicas e psicológicas que distinguem o macho da fêmea permitindo a reprodução sexual e assegurando a continuidade das espécies. Os processos de diferenciação sexual são realizados durante o desenvolvimento embrionário, onde ocorre a proliferação, diferenciação e maturação das células germinativas e primordiais, precursoras (...) de ovócitos e espermatozoides em fêmeas e machos, respectivamente. Assim, os embriões machos e fêmeas iniciam o seu desenvolvimento de forma semelhante, de modo que em ambos os sexos se estabelecem em estruturas idênticas a partir das quais se formarão os órgãos reprodutores correspondentes a cada sexo. O conhecimento da origem e do desenvolvimento do aparelho genital é indispensável para entender sua função e as alterações que produzem infertilidade ou esterilidade. 1.2 DETERMINAÇÃO DO SEXO CROMOSSÔMICO Nos mamíferos, o sexo cromossômico é determinado no momento da fertilização, quando um óvulo, que contém um cromossomo X, é fecundado por um espermatozoide portador de um cromossomo X ou um cromossomo Y. No primeiro caso, o complemento cromossômico seria XX, o que originaria uma fêmea (sexo homogamético), e no segundo daria como resultado um macho com a fórmula cromossômica XY (sexo heterogâmico). 1.3 A GÔNADA INDIFERENCIADA A primeira manifestação das gônadas se aprecia no embrião em forma de um par de eminências longitudinais chamadas cristas ou dobras gonodais, situadas em ambos os lados da linha média entre os mesonefros (rins em desenvolvimento) e do mesentério dorsal. Nos embriões dos mamíferos, as células germinativas primordiais (CGP) manifestam-se em estágios precoces do desenvolvimento, podendo ser detectadas pela primeira vez na metade da gastrulação. As CGP são células grandes, de citoplasma claro e núcleo grande e redondo, localizadas na parede do saco vitelino, perto do alantoide. Essas células possuem grande capacidade de proliferação e vão migrar desde o endoderma do intestino e o epitélio do saco vitelino, através do mesentério, até as cristas gonodais. Isso ocorre por volta do 26° dia da gestação no bovino. Sua migração realiza-se graças aos movimentos de translocação passiva e deslocamento ameboide ativo. Desconhece-se o mecanismo pelo qual estas células são dirigidas para as cristas gonodais, porém foram estudadas algumas moléculas que se expressam durante sua migração e que poderiam desempenhar um papel importante na diferenciação deste tipo celular. A fosfatase alcalina é uma enzima que tem sido usada como marcador de CGP para determinar a sua origem e migração. Num estudo recente, foi inserido um marcador fluorescente que se exprime unicamente nas células germinativas primordiais de embriões transgênicos, e utilizando este marcador e a fosfatase alcalina determinou-se a origem e o padrão de migração destas células. O primeiro sinal de diferenciação das células germinativas primordiais é a expressão de fosfatase alcalina, e esta apareceu pela primeira vez na parte mais posterior da linha primitiva. No sétimo dia de desenvolvimento no embrião do camundongo, o endoderma visceral (AF+) é substituído pelo endoderma definitivo (AF-) originado na parte anterior da linha primitiva. O fator de transcrição Oct-4 é expresso nas CGP de ambos os sexos, pelo que acredita-se estar envolvido na mantença a totipotêncialidade das células. O receptor tirosina quinase, cujo ligante é o fator de Steel, é outra das moléculas que expressam as CGP. Tem sido demonstrado que este receptor possui um papel muito importante na sobrevivência deste tipo celular. Existem outros fatores que promovem a sobrevivência e/ou proliferação de CGP in vitro. Em experiências realizadas com o fator de transformação beta I (TGFβ-I), observou-se que este tem um efeito negativo sobre a proliferação das CGP. Outra atividade que tem sido postulada a este fator é o de um agente quimioatraente que possivelmente possa direcionar a migração destas células para a gônada. a) Um formado pelas células germinativas primordiais (precursoras dos gametas masculinos ou femininos), rodeadas de células somáticas das quais posteriormente se derivarão as células de Sertoli no macho e as células da granulosa na fêmea. b) O tecido que formará o estroma da gônada: tecido conjuntivo, vasos sanguíneos e as células intersticiais com atividade esteroidogênica (células de Leydig no testículo e a teca interna do ovário). As células somáticas do primórdio gonodal originam-se do mesoderma. Inicialmente são de três tipos: mesenquimáticas, mesoteliais e endoteliais. As células mesenquimáticas e mesoteliais iniciam grande atividade proliferativa ao chegar as CGP. Observa-se então uma condensação de células de origem mesotelial e mesenquimatoso que forma um agregado compacto denominado "blastema gonodal". A partir deste primórdio embrionário, diferenciam-se dois tecidos gonodais: os cordões sexuais e o estroma. Os cordões sexuais são arranjos epiteliais que se encontram delimitados por uma folha basal, e dentro deles encontramos as CGP. Por sua vez, no estroma encontram-se células do tipo mesenquimático e vasos sanguíneos. Neste momento, as gônadas são indiferenciadas e bipotencialmente sexuais, sendo impossível diferenciar, morfologicamente, uma gônada masculina de uma feminina, mas no caso dos machos genéticos já existe uma diferenciação da gônada a nível molecular. Nesta fase já se encontram presentes as estruturas das quais se desenvolvem os dutos mesonéfricos ou de Wolff precursores do aparelho genital masculino e os dutos paramesonéfricos ou de Müller que darão origem ao aparelho reprodutor feminino. Há uma série de fatores envolvidos no desenvolvimento precoce da gônada, entre os quais o fator esteroidogênico I (SFI: Steroidogenic fator l), que é um membro da subfamília de receptores nucleares, receptores órfãos. Este fator de transcrição tem um local de ligação ao DNA composto por dois dedos-de-Zinc. O SFI foi identificado como um ativador de genes envolvidos na biossíntese de esteroides em diferentes células. O SFI está presente durante o desenvolvimento embrionário em regiões associadas com funções endócrinas como gônadas, adrenais, pituitárias e hipotálamos. Os animais homozigotos para o gene SFI defeituoso, necessitam de gônadas e adrenais e têm a função gonadotrófica alterada. Os ratos sem SFI carecem de gonadotrofos e têm um desenvolvimento anormal do núcleo ventro-medial do hipotálamo; em particular as gônadas deixam de se desenvolver entre os dias 11 a 15 e degeneram-se por apoptose. No entanto, a crista genital forma-se e é colonizada pelas células germinativas, o que indica que estas continuam a receber o sinal adequado para a sua migração. Portanto, o SFI não está envolvido no desenvolvimento precoce da gônada e do sistema urogenital, mas parece estar envolvido na manutenção do crescimento das células somáticas presentes na gônada indiferenciada. O gene associado ao tumor de Wilms (WTI: Wilm's tumor Associated) está envolvido no desenvolvimento da gônada e do rim. Durante o desenvolvimento embrionário, WTI se expressa em todo o mesoderma intermediário e posteriormente na gônada indiferenciada, bem como no rim em formação. WTI regula o sinal indutivo do mesênquima para o epitélio celômico dos mesonefros. Se este for o caso, então WTI é responsável pelo crescimento da crista genital ao dirigir a entrada do epitélio celômico. Dado que estas células darão origem às células de Sertoli, a carência de WTI pode causar o desenvolvimento de embriões XY como fêmeas simplesmente porque não se formam as células de Sertoli. Em geral, todos os genes importantes na diferenciação do mesoderma intermediário e do sistema urogenital intervêm no desenvolvimento da gônada precoce. 1.4 DIFERENCIAÇÃO GONODAL O desenvolvimento das gônadas e ductos genitais descritos até o momento, é o mesmo para ambos os sexos. Igualmente, os genes descritos, que estão envolvidos no desenvolvimento das gônadas, ductos genitais e migração das células germinativas, afetam igualmente os embriões com genótipo XX ou XY. A gônada primitiva consiste anatomicamente de uma medula (interna) e uma crosta (externa), e de acordo com o local onde ocorre a colonização das células germinativas, diferenciara em testículo ou um ovário, respectivamente. Nos mamíferos, a primeira manifestação estrutural de diferenciação sexual é detectada na gônada dos machos, onde as células germinativas estão localizadas na medula. A diferenciação do testículo inicia-se quando os cordões sexuais se separam do epitélio celômico como consequência dos arranjos produzidos por uma invasão do mesênquima e vasos sanguíneos que provoca a compactação dos cordões, agora denominados cordões testiculares. As células que rodeiam os cordões se achatam e formam as células mioides, que são responsáveis pela formação das membranas basais. As células do epitélio interno, ou seja, as células de Sertoli, têm duas funções principais: o suporte das CGP e a síntese da hormona antimulleriana, responsável pela regressão dos ductos de Müller e secretada durante o período de diferenciação sexual. As células do estroma que rodeiam os cordões testiculares diferenciam-se para formar vários tipos de células: células mioides, fibroblastos, endotélio e células de Leydig, que são as mais importantes pela sua atividade endócrina. Posteriormente, os cordões testiculares dão origem aos túbulos seminíferos, que contêm o epitélio que produzirá os espermatozoides ao chegar à puberdade. Na fêmea, durante os estágios iniciais de diferenciação gonodal, não se observam mudanças em relação à gônada indiferenciada, só pode-se observar um certo crescimento devido à proliferação de células somáticas e germinativas. As células germinativas iniciam um período de proliferação, que termina com o início da meiose. Iniciada a meiose, dá-se o processo de foliculogênese; neste momento os cordões epiteliais se fragmentam, de tal maneira que cada ovócito fica rodeado de células epiteliais cobertas por uma folha basal fina (figura 1). Para que a gônada primitiva se desenvolva em testículo é indispensável a presença do cromossoma Y, independentemente do número de cromossomas X que contenha o genoma de um indivíduo. O gene determinante do testículo encontra-se localizado no cromossoma Y, denominado sry em ratos e SRY em humanos. O gene sry se expressa durante o desenvolvimento embrionário na crista genital de embriões de camundongos. A expressão é detectável no dia 10,5, pouco depois do aparecimento das cristas genitais, atinge o seu máximo durante o dia 11,5 e mantém-se até pouco depois de ocorrerem os primeiros sinais morfológicos de diferenciação testicular no dia 12,5. Este padrão de expressão é compatível com a teoria de que sry atua induzindo a ativação dos genes (figura 2) que conduzem ao desenvolvimento testicular, sem que exista a necessidade da expressão contínua de sry para manter a diferenciação do testículo após o dia 12,5. Como mencionado anteriormente, a gônada primitiva é composta por vários tipos de células. No entanto, as células germinativas primordiais não são o local de expressão do sry, já que os embriões que necessitam de células germinativas mantêm a expressão de sry e desenvolvem o sexo gonodal normalmente. As células somáticas na gônada em desenvolvimento incluem também as células de suporte. Sabe-se que é nestas células que o sry é expresso para que se diferenciem em células de Sertoli, e a expressão transitória de sry indica que deve ativar a outros genes para a manutenção das células de Sertoli. Uma vez diferenciadas as células de Sertoli, elas se encarregarão da diferenciação do resto das células na gônada. -/- Figura 1: Representação da diferenciação dos órgãos genitais internos. Adaptado de BRONSON, 1989. Figura 2: Cascada de genes envolvidos na diferenciação sexual, adaptado de KOOPMAN, 1999. O fator sry é necessário para a diferenciação do testículo. Embora não se conheçam os genes que provavelmente regulam esse gene, estudos realizados em camundongos demonstram que este gene parece coordenar-se com certos genes autossômicos. Entre estes genes autossômicos, o sox9, que é produzido pelas células de Sertoli uma vez que são estimuladas por sry, de modo que sox9 é um dos genes relacionados estruturalmente com sry. O Sox9 funciona como um fator de transcrição, mas não se sabe se a proteína tem qualquer outra função estrutural; este gene exprime-se abundantemente nos condrócitos e está relacionado com defeitos do aparelho ósseo chamados displasia campomélica. Curiosamente, os pacientes XY com esta condição sofrem frequentemente de reversão do sexo. O Sox9 é um dos poucos genes, além do SRY, do qual as mutações demonstraram interferir com a determinação sexual masculina. No entanto, apenas 75% dos pacientes com anomalias esqueléticas de tipo displasia campomélica têm reversão sexual e não foram encontrados casos de reversão sexual devido a um defeito de Sox9 que não seja acompanhado de defeitos esqueléticos. Isso indica que o Sox9 é apenas um membro da rede de genes que são ativados para determinar a diferenciação sexual, enquanto a rota que rege a condrogênese é mais sensível a perturbações deste. O momento em que se detecta a expressão do gene Sox9 (11dpc em ratos) coincide com a máxima expressão de sry, o que poderia indicar a possibilidade de que sry regule positivamente a Sox9. De fato, na região do promotor de Sox9 há um local de união ao que potencialmente se pode unir o sry. A expressão de Sox9 durante a diferenciação sexual sugere um papel abaixo de sry na diferenciação das células de Sertoli. O cromossoma X também é importante na diferenciação gonodal. O gene DAX-I foi isolado do lócus DSS (Dosage sensitive sex reversal) do cromossoma X. DAX-I é parte da cascata de determinação sexual, mas não é necessário para a formação do testículo. DAX-I é um membro dos receptores nucleares conhecidos como receptores órfãos. Este gene demonstrou ser um poderoso repressor da transcrição de SFI e de vários genes. Os padrões de expressão de DAX-I são complementares daqueles de SFI, ambos expressos nas cristas genitais. Em resumo, dada a evidência exposta, desenvolveu-se a hipótese de que DAX-I é um antagonista de sry; esse antagonismo é dependente dos níveis relativos de DAX-I e sry e de um limiar que varia de espécie para espécie. A DAX-I foi classificada como o gene antitestículo. Na fêmea (cariótipo XX) é importante que ocorra a inativação de um dos cromossomas sexuais X para que se mantenha o equilíbrio genético ao igualar o conteúdo de DNA dos cromossomas. Esse cromossoma inativado constitui o chamado corpúsculo de Barr. No entanto, para que a meiose se realize, é necessário dos dois cromossomas X ativos nos ovócitos para assegurar a diferenciação ovárica e a fertilidade. 1.5 DIFERENCIAÇÃO DOS DUCTOS SEXUAIS O embrião possui, além das gônadas indiferenciadas, dois sistemas de ductos: os de potencialidade masculina denominam-se ductos de Wolff ou mesonéfricos, e os de potencialidade feminina se chamam ductos de Müller ou paramesonéfricos (figura 1). Se a diferenciação gonodal levou à formação de um testículo, a partir do ducto mesonéfrico ou de Wolff se desenvolverão os ductos eferentes, o epidídimo, os ductos deferentes e as vesículas seminais. As hormonas importantes no desenvolvimento do aparelho genital masculino são a testosterona, produzida pelas células de Leydig, e sua forma 5α reduzida, a 5α di-hidrotestosterona. Acredita-se que a testosterona é responsável pela virilização dos ductos de Wolff, e a di-hidrotestosterona dos órgãos genitais externos. No macho, os canais de Müller atrofiam-se devido à ação de uma hormona fetal de origem testicular denominada hormona inibidora das estruturas de Müller (HIM) ou hormona antimulleriana. Este processo começa assim que os cordões espermáticos se formam e se diferenciam as células de Sertoli. A existência de HIM foi proposta baseada em estudos realizados em bezerras freemartin, devido à existência de uma hormona responsável pela atresia dos ductos de Müller que na fêmea dá origem ao útero e aos ovidutos. Essa hormona provoca a involução do aparelho genital do bovino nas gestações gemelares nas quais os produtos de diferente sexo têm comunicação sanguínea por ter ocorrido a anastomose dos vasos de ambas as placentas (figura 3). A HIM é uma glicoproteína pertencente à subfamília de TGFβ, é expressa pelas células que darão origem às células de Sertoli e é um dos primeiros marcadores de diferenciação nestas células. A HIM é secretada na vida adulta pelas células de Sertoli no testículo e por células da granulosa no ovário. No rato a HIM é expressa-se no 12° dia em um teste padrão que segue muito de perto o aumento na expressão de sry. No macho, esta secreção de HIM continua durante a vida fetal e adulta, contudo os níveis de HIM declinam na puberdade devido a um aumento na secreção de testosterona. Vários fatores intervêm na regulação do gene de HIM, incluindo os acima descritos SFI e Sox9. O gene HIM contém segmentos de DNA que são conservados em várias espécies de vertebrados. Existe um nexo de ligação para SFI, que ativa a transcrição de HIM. A mutação no local de ligação de SFI resulta em reversão do sexo em indivíduos XY incluindo genitais femininos normais, presença de um útero formado enfatizando a importância de SFI na determinação sexual e na expressão de HIM. Embora SFI seja um bom candidato como regulador de HIM, é expresso em outras células, como as de Leydig e adrenais, que não expressam HIM. Em contrapartida, Sox9 é expresso unicamente nas células de Sertoli que são as produtoras de HIM. O gene HIM também tem um nexo de ligação para Sox9. Além disso, Sox9 pode atuar sinergicamente com SFI para promover a secreção de HIM. Ao contrário destes dois fatores de transcrição, DAX-I antagoniza a ação de Sox9 e provavelmente SFI sobre o promotor de HIM. Assim, para que as células de Sertoli secretem HIM, a transcrição de DAX-I deve diminuir. Figura 3: Representação da diferenciação dos órgãos genitais externos. Adaptado de BRONSON, 1989. Os ductos de Wolff tornam-se o sistema de ejaculação do macho. A porção mais próxima dos testículos dá origem ao epidídimo, a parte central ao ducto deferente e a porção mais distal às vesículas seminais. A próstata e a parte membranosa da uretra do macho desenvolvem-se a partir da porção pélvica do seio urogenital. A virilização e diferenciação dos ductos de Wolff dependem da produção de testosterona pelo testículo. Quanto aos órgãos genitais externos do macho, o tubérculo genital é ampliado e as dobras uretrais se fundem para formar a uretra peniana. A fusão das dobras uretrais aproxima os tubérculos genitais para formar o escroto (figura 4). Figura 4: Diferenciação do aparelho genital da fêmea e do macho. Adaptado de KOOPMAN, 1989. A diferenciação dos órgãos genitais da fêmea ocorre de forma passiva, já que a ausência de testículos e por isso da hormona inibidora dos ductos de Müller (HIM), assim como dos andrógenos virilizantes, favorece o desenvolvimento dos ductos de Müller, enquanto os de Wolff sofrem atrofia. A porção cefálica dos ductos de Müller dá origem aos ovidutos, que na sua terminação caudal se fundem com o útero. O contato dos ductos de Müller com o seio urogenital induz uma intensa proliferação celular que resulta na formação da área uterovaginal localizado entre o seio urogenital e os ductos de Müller. As células do prato uterovaginal proliferam e aumentam a distância entre as duas estruturas criando o espaço que formará a vagina quando o prato é canalizado e forma um lúmen. Em contraste com o que ocorre no macho, na fêmea a maior parte do seio urogenital se mantém exposta na superfície da abertura onde desembocam a vagina e a uretra. O tubérculo urogenital da fêmea tem um crescimento limitado e forma o clitóris. A sequência de passos da diferenciação sexual do aparelho genital é resumida na tabela 1. Tabela 1: Destino em desenvolvimento dos rudimentos sexuais dos fetos macho e fêmea dos mamíferos -/- Rudimento sexual Macho Fêmea Gônada Testículo Ovário Ductos de Müller (Paramesonéfricos) Vestígios Útero, parte da vagina, ovidutos Ductos de Wolff (Mesonéfricos) Ductos eferentes deferentes, epidídimo, vesículas seminais Vestígios Seio urogenital Uretra, próstata, glândulas bulbouretrais Parte da vagina, uretra, vestíbulo, glândulas vestibulares Tubérculo genital Pênis Clitóris Pregas vestibulares Escroto Lábios vulvares Fonte: HAFEZ, 2004. 1.6 DIFERENCIAÇÃO SEXUAL DO HIPOTÁLAMO Os processos de diferenciação sexual não se limitam apenas às células somáticas do organismo do feto, mas incluem também os centros nervosos superiores do cérebro. Assim, da mesma maneira que a gônada e os ductos sexuais se desenvolvem para o tipo feminino ou masculino, propôs-se que o cérebro pode ser "masculinizado" ou permanecer "Feminizado". A diferenciação do hipotálamo vai depender do ambiente esteroidal do neonato e ocorre na fase perinatal. Estes eventos serão de grande transcendência na vida reprodutiva do indivíduo. Tanto a fêmea como o macho nascem com a capacidade de secreção de gonadotropinas de acordo com um padrão cíclico; contudo, no macho, a exposição do hipotálamo à ação dos andrógenos testiculares durante os primeiros dias da vida extrauterina provoca a masculinização, com o qual o hipotálamo do macho é programado para que a secreção de gonadotropinas se realize a um ritmo relativamente constante por parte da hipófise (secreção tônica). Na fêmea, tanto a secreção tônica como a cíclica se conservam. No entanto, observou-se que a injeção de testosterona ou o transplante de testículo na rata fêmea durante os primeiros dias de vida, suprime a sua futura atividade estral (secreção cíclica). Por outro lado, se os ovários forem transplantados para o rato macho normal castrado na idade adulta, o animal não desenvolve qualquer atividade cíclica, mas se os machos transplantados forem castrados ao nascer, o ovário é capaz de efetuar mudanças cíclicas e ovulações. Isto foi demonstrado em roedores, mas não em animais domésticos ou na espécie humana. Portanto, o padrão de secreção de gonadotropinas, seja cíclico ou tônico, não depende da hipófise, mas do hipotálamo e sua correta diferenciação. 1.7 CONCLUSÕES A maioria dos conhecimentos no campo da biologia do desenvolvimento e, muito especificamente, dos processos de diferenciação sexual têm sido originados de estudos relacionados com desordens congênitas, que na sua maioria devem-se a defeitos de genes específicos. A análise detalhada destas desordens permitiu entender alguns mecanismos endócrinos, moleculares e genéticos envolvidos na diferenciação sexual. A identificação do gene sry como determinante do testículo foi uma contribuição crucial e abriu as portas à compreensão dos mecanismos moleculares e celulares relacionados com o desenvolvimento do testículo. Se este gene não estiver presente, é criado um programa genético alternativo para levar a cabo a diferenciação gonodal para o ovário. Finalmente, devemos ter presente que é necessária uma correlação entre mudanças morfológicas e expressão de genes durante o desenvolvimento para entender os mecanismos relacionados com a diferenciação. -/- Apoio -/- Realização -/- REFERÊNCIAS BIBLIOGRÁFICAS ANDERSON, Robert et al. The onset of germ cell migration in the mouse embryo. Mechanisms of development, v. 91, n. 1-2, p. 61-68, 2000. AUSTIN, Colin Russell; SHORT, R. V. Reproduction in Mammals: Volume 1, Germ Cells and Fertilization. Londres: Cambridge University Press, 1972. BRONSON, Franklin H. Mammalian reproductive biology. Chicago: University of Chicago Press, 1989. BUEHR, Mia. The primordial germ cells of mammals: some current perspectives. Experimental cell research, v. 232, n. 2, p. 194-207, 1997. BYSKOV, Anne G. Differentiation of mammalian embryonic gonad. Physiological reviews, v. 66, n. 1, p. 71-117, 1986. CAPEL, Blanche et al. Migration of mesonephric cells into the mammalian gonad depends on Sry. Mechanisms of development, v. 84, n. 1-2, p. 127-131, 1999. CAPEL, Blanche. The battle of the sexes. Mechanisms of development, v. 92, n. 1, p. 89-103, 2000. DERIVAUX, Jules; BARNABÉ, Renato Campanarut. Reprodução dos animais domésticos. Zaragoza: Acribia, 1980. DOMENICE, Sorahia et al. Aspectos moleculares da determinação e diferenciação sexual. Arquivos Brasileiros de Endocrinologia & Metabologia, v. 46, n. 4, p. 433-443, 2002. DONAHOE, Patricia K. et al. Mullerian inhibiting substance activity in bovine fetal, newborn and prepubertal testes. Biology of reproduction, v. 16, n. 2, p. 238-243, 1977. HAFEZ, Elsayed Saad Eldin; HAFEZ, B. Reprodução animal. São Paulo: Manole, 2004. HANLEY, Neil A. et al. Steroidogenic factor 1 (SF-1) is essential for ovarian development and function. Molecular and cellular endocrinology, v. 163, n. 1-2, p. 27-32, 2000. HIORT, Olaf; PAUL-MARTIN, H. The molecular basis of male sexual differentiation. European journal of endocrinology, v. 142, n. 2, p. 101-110, 2000. HOLY, Lubos; MARTÍNEZ JÚSTIZ, G. Colab. Biología de la reproducción bovina. Havana: Revolucionária, 1975. JOSSO, Nathalie et al. The role of anti-Müllerian hormone in gonadal development. Molecular and cellular endocrinology, v. 145, n. 1-2, p. 3-7, 1998. JOST, Alfred et al. Studies on sex differentiation in mammals. In: Proceedings of the 1972 Laurentian Hormone Conference. Londres: Academic Press, 1973. p. 1-41. KNOBIL, Ernst. Knobil and Neill's physiology of reproduction. EUA: Gulf Professional Publishing, 2006. KOFMAN ALFARO, S.; MERCHANT LARIOS, H.; PEREZ PALACIOS, G. Diferenciacion sexual. I. Bases biologicas del dimorfismo sexual. Rev. invest. clín, p. 349-59, 1982. KOOPMAN, Peter. Sry and Sox9: mammalian testis-determining genes. Cellular and Molecular Life Sciences CMLS, v. 55, n. 6-7, p. 839-856, 1999. MCDONALD, L. E. Veterinary endocrinology. Lea & Febiger, Philadelphia, Pa, 1969. MEIZEL, S.; JOHNSON, M. H. Development in mammals. MH Johnson, Ed, v. 3, p. 1-64, 1978. MELLO, Maricilda Palandi de; ASSUMPÇÃO, Juliana de G.; HACKEL, Christine. Genes envolvidos na determinação e diferenciação do sexo. Arquivos Brasileiros de Endocrinologia & Metabologia, v. 49, n. 1, p. 14-25, 2005. -/- REFERÊNCIAS BIBLIOGRÁFICAS MERCHANT-LARIOS, H. Ovarian differentiation. The Vertebrate Ovary, p. 47-81, 1978. MIES FILHO, Antonio. Reprodução dos animais. Porto Alegre: Sulina, 1987. NEF, Serge; PARADA, Luis F. Hormones in male sexual development. Genes & Development, v. 14, n. 24, p. 3075-3086, 2000. PARKER, Keith L.; SCHEDL, Andreas; SCHIMMER, Bernard P. Gene interactions in gonadal development. Annual review of physiology, v. 61, n. 1, p. 417-433, 1999. SWAIN, Amanda; LOVELL-BADGE, Robin. Mammalian sex determination: a molecular drama. Genes & development, v. 13, n. 7, p. 755-767, 1999. WILHELM, Dagmar; PALMER, Stephen; KOOPMAN, Peter. Sex determination and gonadal development in mammals. Physiological reviews, v. 87, n. 1, p. 1-28, 2007. WILSON, Jean D.; GRIFFIN, James E.; GEORGE, Fredrick W. Sexual differentiation: early hormone synthesis and action. Biology of reproduction, v. 22, n. 1, p. 9-17, 1980. -/- Emanuel Isaque Cordeiro da Silva Belo Jardim, 07 de Maio de 2020. (shrink)

Abstract: Introspection reveals that one is frequently conscious of some form of inner speech, which may appear either in a condensed or expanded form. It has been claimed that this speech reflects the way in which language is involved in conscious thought, fulfilling a number of cognitive functions. We criticize three theories that address this issue: Bermúdez’s view of language as a generator of second-order thoughts, Prinz’s development of Jackendoff’s intermediate-level theory of consciousness, and Carruthers’s theory of inner speech as (...) a rehearsal of action-schemata. We contend they have problems to account for those cases in which inner speech is fragmentary, and for the difference with those instances in which it appears as more sentence-like. In addition, we present verbal overshadowing as a phenomenon that neither of them can easily explain. Finally, we propose an account in which inner speech is fundamentally silent outer speech and argue that it is more explanatory than the alternatives. (shrink)

Philosophical discussions on cognition (ranging from computational and representational approaches to the 4E fra-mework), on the one hand, and those on sound and auditory per-ception, on the other, have hitherto remained apart. In this paper, the author addresses the concept of “auditory cognition”. While committing to the conceptual analysis of the latter, the author in-troduces the discussion of modality-specif ic approaches. Since the contributions made so far in the domain of the philosophy of sound and auditory perception are prone to (...) favor computational-repre-sentational approaches —e.g., the perceptual coherence model (Méndez-Martínez, 2022)—, the exploration of 4E possibilities will lay down a series of questions aimed to encourage future developers from that viewpoint. In that sense, the author explores conceptual entailments and connections between the genealogy of theories within the philosophy of sound and auditory perception, and the hypothesis of the extended cognition (EX). (shrink)

(This is the prologue to a book examining the causes of day-to-day strife in the populations of modern democracies vying to live and assert the freedoms promised to them by systems of governance supposed and expected to represent them.) -/- Voters, the population at large, no longer want representation but participation. But how is that achieved? The technological means are certainly there for each one of us to express opinion, bring forth ideas, mobilise groups with common purpose. -/- The growth (...) of the social, political, and professional consciousness has gone beyond the stage of conformism and acceptance. The time is ripe for mechanisms that create profound bottlenecks to be superseded by better means, less procedure, less posturing, full involvement at grass roots. (shrink)

(This is one of the essays to be included in a book examining the causes of day-to-day strife in the populations of modern democracies vying to live and assert the freedoms promised to them by systems of governance supposed and expected to represent them.) -/- The emigrant / immigrant / migrant makes a conscious, relatively difficult decision to exchange what s/he knows for what is not known at all but in promise. The choice is often stark and carries with it (...) the imposition of restrictions on what will be possible in terms of existing customs, culture, and direct involvement with friends and family, and signifies a leap of faith into what appears to be a better, more prosperous, happier future. -/- At times, such choice is unavoidable, others it turns out to be a gamble of sorts, but, no matter what the cause, reason, or desire underlying an act of migration away from what the emigrant has known, it never turns out to be what was expected and no amount of preparation allows the emigrant to anticipate what will in fact occur. Migration is the founding of a new life in all the aspects of the emigrant’s existence. (shrink)

Intentionalism is the view that perceptual phenomenology depends on perceptual content. The aim of this paper is to make explicit an ambiguity in usual formulations of intentionalism, and to argue in favor of one way to disambiguate it. It concerns whether perceptual phenomenology depends on the content of one and only one representation (often construed as being identical to a certain perceptual experience), or instead depends on a collection of many different representations throughout the perceptual system. We argue in favor (...) of the latter option. Intentionalism so conceived can make better sense of contemporary neuroscience of perception, and is better equipped to confront several influential objections to traditional intentionalism. (shrink)

Entre las virtudes de la explicación de Gewirth, destaca su potencial para delimitar el objeto de los derechos humanos y contener la expansión incontrolada del discurso de los derechos. Al ceñirse al contexto de la acción y a los rasgos necesarios para la agencia intencional, dicha explicación proporciona un fundamento basado en la autonomía y libertad de los agentes intencionales prospectivos, tan solo limitadas por aquellas exigencias racionales derivadas de la igual consideración del status moral de todos los demás agentes (...) intencionales prospectivos quienes, en última instancia, se identifican con la humanidad en su conjunto mediante las ideas implícitas de reciprocidad y responsabilidad moral por los actos propios. En definitiva, el argumento de Gewirth desemboca en el PCG para proporcionar un fundamento racional de los derechos humanos limitado a la estructura normativa de la acción, la agencia intencional prospectiva potencial o actual de todos los seres humanos y los bienes necesarios para la acción intencional: la libertad y el bienestar. (shrink)

We aim to map out the points of confluence between phronesis and emotion, as well as the nature of this confluence. We do so based on philosophical and psychological explanations of emotions and phronesis. Making sound decisions, which requires phronesis, is an important matter, but its relationship with emotions has only just begun to be studied. We propose that the interplay between phronesis and emotion is possible (rather than inevitable) because both have a cognitive-behavioural structure and because emotions are hierarchical. (...) In the case of emotion, there are no gaps between the cognitive and behavioural dimensions, while phronetic acts can only manifest if cognitive and behavioural aspects are activated simultaneously. The confluence we identify is bidirectional and involves mutual enrichment in this dual dimension. On the one hand, phronesis enriches emotion because it involves extra subjective knowledge and allows emotion to be directed towards higher goals. On the other hand, emotions enrich phronesis because phronesis makes decisions based on sensitive preferences and because emotions confirm correctness in the context of moral decision making. Methodologically, this study is theoretical and interdisciplinary, as our approach is both philosophical and psychological. Philosophy helps us to improve the conceptual foundation of the construct, while psychology provides an analysis of the emotional process, allowing philosophy to understand how emotional evaluation-assessment and motivation lead to decision making. (shrink)

(This is one of the essays to be included in a book examining the causes of day-to-day strife in the populations of modern democracies vying to live and assert the freedoms promised to them by systems of governance supposed and expected to represent them.) "The artisan of old, the artist, the researcher, the developer, and the scientist today have this in common, that in refining, perfecting and pushing the boundaries of their respective crafts, they cannot achieve satisfaction or adequately perform (...) without simultaneously and progressively adapting, re-configuring, developing, and continuously changing the techniques and approaches they employ together with the tools and parameters for gauging actual success within the realm that only rigorous acquaintance, imaginative freedom, cross-functional, multifaceted learning, and distributive interaction may allow.". (shrink)

An important objection to signaling approaches to representation is that, if signaling behavior is driven by the maximization of usefulness, then signals will typically carry much more information about agent-dependent usefulness than about objective features of the world. This sort of considerations are sometimes taken to provide support for an anti-realist stance on representation itself. The author examines the game-theoretic version of this skeptical line of argument developed by Donald Hoffman and his colleagues. It is shown that their argument only (...) works under an extremely impoverished picture of the informational connections that hold between agent and world. In particular, it only works for cue-driven agents, in Kim Sterelny’s sense. In cases in which the agents’ understanding of what is useful results from combining pieces of information that reach them in different ways, and that complement one another, maximizing usefulness involves construing first a picture of agent-independent, objective matters of fact. (shrink)

There has been much discussion of so-called teleosemantic approaches to the naturalization of content. Such discussion, though, has been largely confined to simple, innate mental states with contents such as ?There is a fly here.? Even assuming we can solve the issues that crop up at this stage, an account of the content of human mental states will not get too far without an account of productivity: the ability to entertain indefinitely many thoughts. The best-known teleosemantic theory, Millikan's biosemantics, offers (...) an account of productivity in thought. This paper raises a basic worry about this account: that the use of mapping functions in the theory is unacceptable from a naturalistic point of view. (shrink)

In the predominant pedagogical frameworks of our days, rationalist, voluntarist or sentimentalist reductionisms are seen, from which it is about educating people regardless of their Origin and the singular and unrepeatable originality with which they have been given to existence. Faced with this anthropocentric confinement, Sensitive Education arises so that every person, regardless of their culture and creed, remains sensitive to their Origin and captures their own originality, which in the end is what they must accept and try to manifest (...) with education. When educational systems are not sensitive to the original, they develop falsehood. With the aim of consolidating the validity of the pedagogical-anthropological foundation of Sensitive Education, this study seeks to show how Edith Stein’s argument explains Sensitive Education, although Stein, after her conversion, uses explicitly Christian terminology. Stein’s direct terms serve to describe Education Sensitive to the Holy Spirit, but for those who do not start from a Christian worldview, Education Sensitive to Origin can be a more non-denominational way of approaching the Original. To achieve this objective, an analytical-synthetic methodology has been followed in which essential features of Sensitive Education are proposed and in a hermeneutic manner, arguments are extracted from the writings of Edith Stein that offer support for these essential features. In conclusion, it is sufficiently shown that Edith Stein’s thought provides a pedagogical foundation to Sensitive Education. (shrink)

Categorization behavior can be fruitfully analyzed in terms of the trade‐off between as high as possible faithfulness in the transmission of information about samples of the classes to be categorized, and as low as possible transmission costs for that same information. The kinds of categorization behaviors we associate with conceptual atoms, prototypes, and exemplars emerge naturally as a result of this trade‐off, in the presence of certain natural constraints on the probabilistic distribution of samples, and the ways in which we (...) measure faithfulness. Beyond the general structure of categorization in these circumstances, the same information‐centered perspective can shed light on other, more concrete properties of human categorization performance, such as the results of certain prominent experiments on supervised categorization. (shrink)

This paper addresses the relationship between logic and phenomenology at a historical moment that precedes the big divide between analytic philosophy and phenomenology. In analysing alternative derivations of phenomenological logic, the discussion focuses on the case of two notorious neo-Kantian Mexican philosophers from the first half of the XXth century: Adalberto García de Mendoza and Francisco Larroyo. It is argued that both García de Mendoza and Larroyo made an original contribution to the discussion on the relationship between phenomenology and logic. (...) Further, this paper attempts to present a charitable portrait of both García de Mendoza and Larroyo in contrast to underrating approaches to both Mexican philosophers. In so doing, this paper adopts a historical—though not strictly biographical— perspective. While it is true that neither García de Mendoza nor Larroyo could be considered “phenomenologists” or “Husserl scholars” according to contemporary standards it is worth noticing that they engage in the discussion of phenomenological logic. Both authors had the project of developing a grounding logic for science and, as the article argues, both attempts could be even linked. Their project, however, was unfinished. The plan of this paper is as follows. The first section addresses the problem of the relationship between logic and phenomenology. The second one focuses on García de Mendoza’s contributions. The third one elaborates on Larroyo’s Logic of Sciences. And, finally, the fourth section elaborates on these Mexican philosophers’ contributions to education, making as well further observations concerning the similarities and differences between both authors. (shrink)

Anydomainofscientificresearchhasitssustainingorthodoxy. Thatis, research on a problem, whether in astronomy, physics, or biology, is con- ducted against a backdrop of broadly shared assumptions. It is these as- sumptionsthatguideinquiryandprovidethecanonofwhatisreasonable-- of what "makes sense." And it is these shared assumptions that constitute a framework for the interpretation of research results. Research on the problem of how we see is likewise sustained by broadly shared assump- tions, where the current orthodoxy embraces the very general idea that the business of the visual system is to (...) create a detailed replica of the visual world, and that it accomplishes its business via hierarchical organization and by operatingessentiallyindependently of other sensorymodalitiesas well as independently of previous learning, goals, motor planning, and motor execution. (shrink)

We explore the contribution made by oscillatory, synchronous neural activity to representation in the brain. We closely examine six prominent examples of brain function in which neural oscillations play a central role, and identify two levels of involvement that these oscillations take in the emergence of representations: enabling (when oscillations help to establish a communication channel between sender and receiver, or are causally involved in triggering a representation) and properly representational (when oscillations are a constitutive part of the representation). We (...) show that even an idealized informational sender–receiver account of representation makes the representational status of oscillations a non-trivial matter, which depends on rather minute empirical details. (shrink)

I explore some of the connections between Grahek's model of asymbolic pain, as developed in Feeling Pain and Being in Pain, and the contemporary intentionalist discussion over evaluativist and imperativist models of pain. I will sketch a Grahekian version of imperativism that is both true to his main insights and better at confronting some of the challenges that his theory has faced since its publication.

Recent work on signaling has mostly focused on communication between organisms. The Lewis–Skyrms framework should be equally applicable to intra-organismic signaling. We present a Lewis–Skyrms signaling-game model of painful signaling, and use it to argue that the content of pain is predominantly imperative. We address several objections to the account, concluding that our model gives a productive framework within which to consider internal signaling.

I respond to an objection recently formulated by Barlassina and Hayward against first-order imperativism about pain, according to which it cannot account for the self-directed motivational force of pain. I am going to agree with them: it cannot. This is because pain does not have self-directed motivational force. I will argue that the alternative view—that pain is about dealing with extramental, bodily threats, not about dealing with itself—makes better sense of introspection, and of empirical research on pain avoidance. Also, a (...) naturalistic theory of body-involving commands falls straightforwardly out of our most prominent naturalistic metasemantic accounts, while the token-reflexive contents that would underlie self-directed motivation are more problematic. (shrink)

El autor del artículo edita y estudia una inscripción funeraria inédita de época helenística encontrada en Aptera (Creta) por la arqueóloga griega V. Ninioú-Kindelí. El texto de la inscripción dice así: A) Σώσανδρος | Βίτωνος. B) Ἀμφιμήδης | Σωσάνδρω.

The changes known as the loss of inflexions in English (11th- 15th centuries, included) were prompted with the introduction of a new mode of thinking. The mode of thinking, for the Anglo-Saxons, was a dynamic way of conceiving of things. Things were considered events happening. With the contacts of Anglo-Saxons with, first, the Romano-British; second, the introduction of Christianity; and finally with the Norman invasion, their dynamic way of thinking was confronted with the static conception of things coming from the (...) Mediterranean. The history of English from the 11th to the 15th century meant the introduction, confrontation and adoption of a new mental conception of things, the static way of conceiving of things, both modes of thinking defining the language today. (shrink)

This paper provides a semantic analysis of the particles afinal (European Portuguese) and alla fine (Italian) in terms of the notion of truth unpersistence, which can be situated at the intersection of epistemic modality and discourse structure. In the analysis proposed, the particles are propositional operators and require that the truth of a proposition p* fail to persist through a temporal succession of epistemic states, this proposition being incompatible with the prejacent, and that the interlocutors share knowledge of a previous (...) epistemic attitude toward p*. We analyze two main cases (plan-related and non plan-related propositions) and also show that these particles are indexical to one (or more) epistemic agent(s) and allow for shifts in perspective. (shrink)

Altruism and solidarity are concepts that are closely related to organ donation for transplantation. On the one hand, they are typically used for encouraging people to donate. On the other hand, they also underpin the regulations in force in each country to different extents. They are often used indistinctly and equivocally, despite the different ethical implications of each concept. This paper aims to clarify to what extent we can speak of altruism and solidarity in the predominant models of organ donation. (...) It also raises the ethical question of whether these categories are adequate as a basis for such models, bearing in mind that organs are a scarce resource and that a shortage of them may mean that fewer lives are saved or improved. (shrink)

It is widely held that it is unhelpful to model our epistemic access to modal facts on the basis of perception, and postulate the existence of a bodily mechanism attuned to modal features of the world. In this paper I defend modalizing mechanisms. I present and discuss a decision-theoretic model in which agents with severely limited cognitive abilities, at the end of an evolutionary process, have states which encode substantial information about the probabilities with which the outcomes of a certain (...) Bernoulli process occur. Thus, in the model, a process driven by very simple, thoroughly naturalistic mechanisms eventuates in modal sensitivity. (shrink)

Despite considerable growth in understanding of various aspects of sporting and exercise embodiment over the last decade, in-depth investigations of embodied affectual experiences in running remain limited. Furthermore, within the corpus of literature investigating pleasure and the hedonic dimension in running, much of this research has focused on experiences of pleasure in relation to performance and achievement, or on specific affective states, such as enjoyment, derived after completing a run. We directly address this gap in the qualitative literature on sporting (...) and exercise embodiment by contributing novel insights on the mind-body pleasures of running via focusing analytic attention towards the pleasures recalled by runners as experienced during positive, rewarding running experiences. Applying conceptual insights drawn from sociological phenomenology, we analyse data from an in-depth, event-focused interview study with distance runners who reported positive, rewarding experiences in recent recreational runs. Through reflexive thematic analysis, we present findings in relation to three themes: (1) ‘running feels like it should’; (2) sensory engagements; and (3) time out. The study contributes fresh perspectives, both conceptually and in relation to data-collection approach, to a small literature on the lived experience of pleasure in sport, exercise and physical cultures. (shrink)

Subjective rights enjoy limited import in Kelsenian theory for whereas the concept of duty underlies every legal norm, that of rights is merely possible and only emerges when the imposition of the sanction attached to the breach of the duty is made dependent upon a subject's will to bring legal action. The presence of secondary norms establishing certain duties of medical professionals on informed consent displays the existence of correlative reflex rights of patients. Yet, together with secondary norms, Western legal (...) systems typically institute norms of competence enabling patients to take legal action when those duties correlative to their reflex right to informed consent have been infringed. Thus, informed consent constitutes a set of subjective legal positions of patients and medical professionals. Despite its excessive formalism and questionable neutrality, the Kelsenian approach permits to define and clearly distinguish patients' genuine rights from mere aspirations devoid of legal basis. (shrink)

Why do we have social norms—of fairness, cooperation, trust, property, or gender? Modern-day Humeans, as I call them, believe these norms are best accounted for in cultural evolutionary terms, as adaptive solutions to recurrent problems of social interaction. In this paper, I discuss a challenge to this “Humean Program.” Social norms involve widespread behaviors, but also distinctive psychological attitudes and dispositions. According to the challenge, Humean accounts of norms leave their psychological side unexplained. They explain, say, why we share equally, (...) but not why we disapprove of those who don’t. I defend the Humean Program against this challenge. In particular, I suggest an idea for how to extend the Program to account for the psychological side of norms. Socially adaptive behaviors aren’t just likely to emerge in a group; They are also likely to be widely taught within it. The transmission of these behaviors through instruction explains why they are associated with distinctive normative attitudes and dispositions. These attitudes play a pedagogical role in helping transmit these behaviors to children and newcomers. (shrink)

Imperativism is the view that the phenomenal character of the affective component of pains, orgasms, and pleasant or unpleasant sensory experience depends on their imperative intentional content. In this paper I canvass an imperativist treatment of pains as reason-conferring states.

In the realm of the philosophy of sounds and auditory experience there is an ongoing discussion concerned with the nature of sounds. One of the contestant views within this ontology of sound is that of the Property View, which holds that sounds are properties of the sounding objects. A way of developing this view is through the idea of dispositionalism, namely, by sustaining the theory according to which sounds are dispositional properties (Pasnau 1999; Kulvicki 2008; Roberts 2017). That portrayal, however, (...) is not sufficient, as it has not inquired the metaphysical debates about dispositions beyond the conditional analysis. In this paper, I try to advance this view by including recent developments (for instance Bird 2007; Vetter 2015) in the field of dispositionalism and I analyse whether this new version can sort out known and new objections to Property View. (shrink)

Our limited a priori-reasoning skills open a gap between our finding a proposition conceivable and its metaphysical possibility. A prominent strategy for closing this gap is the postulation of ideal conceivers, who suffer from no such limitations. In this paper I argue that, under many, maybe all, plausible unpackings of the notion of ideal conceiver, it is false that ideal negative conceivability entails possibility.

Contextualist theorists have recently defended the views (a) that metaphor-processing can be treated on a par with other meaning changes, such as narrowing or transfer, and (b) that metaphorical contents enter into “what is said” by an utterance. We do not dispute claim (a) but consider that claim (b) is problematic. Contextualist theorists seem to leave in the hands of context the explanation about why it is that some meaning changes are directly processed, and thus plausibly form part of “what (...) is said”, while some others are not. While granting the role of context in this respect, we contend that there are that there are elements that play an instrumental role in providing direct access to the metaphorical content, namely, the conventionality of the expressions and the salience of the concepts involved. We will start by criticizing Recanati’s and Relevance Theory’s accounts of metaphor. Then we examine the claims of Carston’s and Giora’s two-process accounts that set the stage for a revision of the main elements involved, namely, the properties of conventionality and salience. Finally we examine a number of representative examples, explaining why some cases involve a direct access to the metaphorical content and others require an intermediate non-figurative interpretation. (shrink)

Reformation commentators were well aware of the allegorical referents for Leviathan and Behemoth in the book of Job, representing the powerful states of Ancient Egypt and Assyria, but played them down. Hobbes did not.

Quentin Skinner’s dedication to investigating Hobbes’s concept of liberty in a number of essays and books has born some unusual fruit. Not only do we see the enormous problems that Hobbes set himself by proceeding as he did, but Skinner’s careful analysis allows us to chart Hobbes’ ingenuity as he tried to steer a path between the Charybdis of determinism and the Scylla of voluntarism – not very successfully, as we shall see. The upshot is a theory of individual freedom (...) and civil liberty to challenge the classical republican traditionThis Article does not have an abstract. (shrink)

This paper brings new work to bear on the perennial question about Hobbes's atheism to show that as a debate about scepticism it is falsely framed. Hobbes, like fellow members of the Mersenne circle, Descartes and Gassendi, was no sceptic, but rather concerned to rescue physics and metaphysics from radical scepticism by exploring corporealism. In his early letter of November 1640, Hobbes had issued a provocative challenge to Descartes to abandon metaphysical dualism and subscribe to a ?corporeal God?; a provocation (...) to which the Frenchman angrily responded, but was perhaps importantly influenced. Hobbes's minimal realism was consonant with atheism, to which Descartes felt he was being forced. Moreover, Hobbes was unrelenting in his battle against Cartesian dualism, for which he saw Robert Boyle's experimental science as a surrogate. (shrink)

This paper addresses the problem of sound individuation (SI) and its connection to sound ontology (SO). It is argued that the problems of SI, such as aspatiality, extreme individuation, indexical perplexity and duration puzzles are due to SO’s uncertainties. Besides, I describe the views in SO, including the wave view (WV), the property view (PV), and the event view (EV), as Casey O’Callaghan defends it. According to O’Callaghan, EV offers clear standards to individuate sounds. However, this claim is countered by (...) the consideration that any view could also defend the standards in SO, and thus, EV does not solve any of the problems mentioned above. As a way of showing the difficulties inherited by sound’s inner ontology, the problem of its linguistic representation is also addressed. The problem of SI can be developed within the frame of the philosophy of language and, specifically, regarding the discussion about mass vs count-­sortal terms. Is the term sound a mass or a count­-sortal? It is shown that, for reasons pertaining SO, the decision regarding the case of sound as a mass or count-­sortal term remains open. SI is, thus, covered from the SO to the philosophy of language. (shrink)

ABSTRACT: Hobbes belonged to philosophical and scientific circles grappling with the big question at the dawn of modern physics: materialism and its consequences for morality. ‘Matter in motion’ may be a core principle of this materialism but it is certainly inadequate to capture the whole project. In wave after wave of this debate the Epicurean view of a fully determined universe governed by natural laws, that nevertheless allows to humans a sphere of libertas, but does not require a creator god (...) or teleology to explain it, comes up against monotheism and its insistence on the incoherence of an ordered world in the absence of a God and his purposes. The following questions were central to this debate: Can we understand the universe as law-governed in the absence of a god? If so, what room is there in a fully determined mechanical universe for human freedom? If humans do enjoy freedom, does the same hold for other animals? Is this freedom compatible with standard views of morality? (shrink)

In the first part of the paper, I present a framework for the description and evaluation of teleosemantic theories of intentionality, and use it to argue that several different objections to these theories (the various indeterminacy and adequacy problems) are, in a certain precise sense, manifestations of the same underlying issue. I then use the framework to show that Millikan's biosemantics, her own recent declarations to the contrary notwithtanding, presents indeterminacy. In the second part, I develop a novel teleosemantic proposal (...) which makes progress in the treatment of this family of problems. I describe a procedure to derive a (unique) homeostatic property cluster [HPC] from facts having to do with the properties that a certain indicator relied on, in the events leading to its fixation in a certain population. This HPC is the one that should figure in the content attribution to the indicator in question. (shrink)

La lengua española nació cuando los hablantes del latín de la península Ibérica cambiaron de modo de pensar. Desde el pensar absoluto latino, asimilaron el pensar substantivo de los griegos a través del pensar cristiano. Haciendo una síntesis de estos tres pensares absolutos, crearon el pensar individual, el pensar abstracto y el pensar real. De este modo crearon lo que se puede designar como una ontología lingüística, la manifestación directa del pensar español. Los modos de pensar tienen que ver con (...) los modos de concebir lo que nos rodea. Ambos a dos preceden y determinan el pensamiento y la manifestación de las actividades libres del hombre, tanto si este es considerado individualmente, como si es considerado socialmente dentro del conjunto de hablantes que participan y contribuyen dentro de una comunidad lingüística. (shrink)

Linguistics of Saying is to be analyzed in the speech act conceived as an act of knowing. The speaking, saying and knowing subject, based on contexts and the principles of congruency and trust in the speech of other speakers, will create meanings and interpret the sense of utterances supplying the deficiencies of language by means of the intellective operations mentally executed in the act of speech. In the intellective operations you can see three steps or processes: first the starting point, (...) intuition or aísthesis; second, the process of abstraction; and third, the inverse: the process of determination or fixing the construct created. (shrink)

Linguistics of saying studies language in its birth. Language is the mental activity executed by speaking subjects. Linguistics of saying consists in analyzing speech acts as the result of an act of knowing. Speaking subjects speak because they have something to say. Tthey say because they define themselves before the circumstance they are in. And this is possible because they are able to know. Speaking, then, is speaking, saying and knowing. In this sense there is a progressive determination. Knowing makes (...) possible saying, and saying determines speaking, or, in other words: speaking involves saying and knowing, and saying involves knowing. The problem thus is to determine the meaningful intentional purpose of the individual speaker to say something in every speech act. (shrink)

“Conciencia”, la sección inicial de la Fenomenología de Hegel, se ocupa de la comprensión del mundo físico que nos rodea. La sección siguiente, “Autoconciencia”, comienza a considerar nuestra comprensión de nosotros mismos y de los otros. Este orden de discusión no es ni arbitrario ni meramente conveniente. Por el contrario, una de las principales lecciones que hemos de aprender hacia el final del desarrollo de la “conciencia” es que nuestra mejor concepción del mundo que es el objeto de nuestras actividades (...) cognitivas es inteligible solo como una parte de un relato que además considera la naturaleza del sujeto que se dedica a tales actividades. La razón para esta transición expositiva es un aspecto importante en el idealismo de Hegel. En este capítulo ofrezco una reconstrucción racional de un argumento que veo como el soporte de esta transición y del tipo de idealismo que esta implica. (shrink)

Some mental states have valence—they are pleasant or unpleasant. According to imperativism, valence depends on imperative content, while evaluativism tells us that it depends on evaluative content. We argue that if one considers valence’s informational profile, it becomes evident that imperativism is superior to evaluativism. More precisely, we show that if one applies the best available metasemantics to the role played by (un)pleasant mental states in our cognitive economy, then these states turn out to have imperative rather than evaluative content, (...) since: (i) they are much more informative about behaviour than they are about the world; and (ii) they occupy a stage in the information-processing chain that is closer to behaviour production than it is to the uptake of sensory information. This is our metasemantic argument for imperativism. (shrink)

Thomas Hobbes's 'Historia Ecclesiastica' presents his views on religion and aims to divert the attention of the public from charges against his being a heretic to placing heresy in pagan history, claiming that Greek philosophers were responsible for introducing heresy in the Christian Church. His book reveals his interest in religious history and the growth of hermeticism and Cabalism in England in his age.

Hegel's interpretation of Sophocles' play Antigone is central to an understanding of woman's role in the Hegelian system. Hegel is fascinated by this play and uses it in both the Phenomenology and the Philosophy of Right to demonstrate that familial ethical life is woman's unique responsibility. Antigone is revealed as the paradigmatic figure of womanhood and family life in both the ancient and modern worlds, although there are fundamental differences between these two worlds for Hegel. Through an immanent critique of (...) both the Phenomenology and the Philosophy of Right which focuses on the role of woman as presented by Hegel in the figure of Antigone, my analysis reveals the limitations of Hegel’s dialectical theory. (shrink)

El lenguaje o la actividad cognoscitiva del ser humano que se debate en su lucha contra la circunstancia en la que le ha tocado vivir es hablar, decir y conocer. El hombre habla porque tiene algo que decir, dice porque se define a sí mismo ante la circunstancia en la que vive en cada momento, y esto es posible porque conoce de forma creativa. En este sentido el decir determina el hablar, por arriba, y el conocer por abajo. El conocer (...) humano empieza en una sensación sensible, la aísthesis, la cual es transformada en un universal, extracto o concepto, logos, el cual es el fruto de una serie de operaciones intelectivas realizadas por el propio sujeto en u solo acto, el acto único de hablar, decir y conocer. (shrink)

The human subject in as much as he knows transforms the sensitive and concrete (the thing perceived) into abstract (an image of the thing perceived), the abstract into an idea (imaginative representation of the thing abstracted), and ideas into contents of conscience (meanings). The last step in the creation of meanings, something being executed in the speech act, consists in fixing the construct mentally created thus making it objectified meanings in the conscience of speakers. The interchange amongst the different steps (...) in the creation of meaning manifests lógos, the state lived by speakers in their interior when speaking, created and developed in words and because of words. (shrink)

Introduction: Traditionally, empathy has been studied from two main perspectives: the theory-theory approach and the simulation theory approach. These theories claim that social emotions are fundamentally constituted by mind states in the brain. In contrast, classical phenomenology and recent research based on enactive theories consider empathy as the basic process of contacting others’ emotional experiences through direct bodily perception and sensation. Objective: This study aims to enrich knowledge of the empathic experience of pain by using an experimental phenomenological method. Method: (...) Implementing an experimental paradigm used in affective neuroscience, we exposed 28 healthy adults to a video of sportspersons suffering physical accidents while practicing extreme sports. Immediately after watching the video, each participant underwent a phenomenological interview to gather data on embodied, multi-layered dimensions (bodily sensations, emotions, and motivations) and temporal aspects of empathic experience. We also performed quantitative analyses of the phenomenological categories. Results: Experiential access to the other person’s painful experience involves four main-themes. Bodily resonance: participants felt a multiplicity of bodily, affective, and kinesthetic sensations. Attentional focus: some participants centered their attention more on their own personal discomfort and sensations of rejection, while others on the pain and suffering experienced by the sportspersons. Kinesthetic motivation: some participants experienced the feeling in their bodies to avoid or escape from watching the video, while others experienced the need to help the sportspersons avoid suffering any injury while practicing extreme sports. Temporality of experience: participants witnessed temporal fluctuations in their experiences, bringing intensity changes in their bodily resonance, attentional focus, and kinesthetic motivation. Finally, two experiential structures were found: one structure is self-centered empathic experience, characterized by bodily resonance, attentional focus centered on the participant’s own experience of seeing the sportsperson suffering, and self-protective kinesthetic motivation; the other structure is other-centered empathic experience, characterized by bodily resonance, attentional focus centered on the sportsperson, and prosocial kinesthetic motivation to help them. Discussion: We show how phenomenological data may contribute to comprehending empathy for pain in social neuroscience. In addition, we address the phenomenological aspect of the enactive approach to the three dimensions of embodiment of human consciousness, especially the intersubjective dimension. Also, based on our results, we suggest an extension of the enactive theory for non-interactive social experience. (shrink)

Both Franz Brentano and Edmund Husserl addressed sound while trying to explain the inner consciousness of time and gave to it the status of a supporting example. Although their inquiries were not aimed at clarifying in detail the nature of the auditory experience or sounds themselves, they made some interesting observations that can contribute to the current philosophical discussion on sounds. On the other hand, in analytic philosophy, while inquiring the nature of sounds, their location, auditory experience or the audible (...) qualities and so on, the representatives of that trend of thought have remained silent about the depiction of sound and the auditory phenomena in the phenomenological tradition. The paper’s intention is to relate both endeavours, yet the perspective carried out is that of analytic philosophy and, thus, I pay special attention to conceptual analysis as a methodological framework. In this sense, I first explain what sound ontology is in the context of analytic philosophy and the views that it encompasses— namely, the Property View (PV), the Wave View (WV) and the Event View (EV)—. Secondly, I address the problems it entails, emphasising that of sound individuation. In a third section, I propose the possibly controversial conjunction of a “Brentano-Husserl Analysis of the Consciousness of Time” (for short “Brentano-Husserl analysis”) and outline the commonalities of both authors, without ignoring its discrepancies. My main focus is Husserl’s 1905 Vorlesungen zur Phänomenologie des Inneren Zeitbewusstseins. While addressing the Brentano-Husserl analysis, I elaborate on the problem of temporal and spatial extension (Raumlichkeit and Zeitlichkeit, respectively) of both consciousness and sound. Such comparison is a key one, since after these two developments, one can notice some theoretical movements concerning the shift of attention from sounds to the unity of consciousness, and how they mirror each other. After examining the controversial claims concerning the temporal and spatial extension of both consciousness and sound, I argue in the concluding paragraphs that while considering the accounts of sound ontology, the Brentano-Husserl analysis would probably endorse a Property View and that this could have interesting consequences for the issue of Sound Individuation. (shrink)