TRANSPORTE DE GAMETAS, FERTILIZAÇÃO E SEGMENTAÇÃO -/- • _____OBJETIVO -/- O entendimento do desenvolvimento embrionário nos estágios iniciais, desde a deposição dos espermatozoides na fêmea, passando pela fertilização deste no ovócito e na formação do zigoto, é de suma importância para diferenciar especialistas em reprodução e manejo reprodutivo no mercado de trabalho e, também, durante a vida acadêmica. Compreender os processos que levam à formação do zigoto na fêmea é essencial para avaliar a capacidade reprodutiva dos animais e, mediante técnicas, (...) avaliar o trato reprodutivo da fêmea e o desenvolvimento do embrião até a formação do feto e, por fim, em um novo animal sadio na propriedade. Com esse trabalho, o estudante de veterinária ou zootecnia, que deseja se aprofundar na fisiologia da reprodução animal, identificará os mecanismos que são envolvidos no transporte dos gametas, bem como os processos que estes últimos devem completar para alcançar a fertilização e para desencadear as divisões iniciais do embrião. -/- • _____INTRODUÇÃO -/- A fertilização ou fecundação é o processo pelo qual os gametas masculino (espermatozoide) e feminino (ovócito) se fundem para gerar um novo indivíduo. Seu êxito depende da culminação adequada dos diferentes processos que devem sofrer os gametas durante sua maduração e percurso; do transporte oportuno destes no trato reprodutivo da fêmea, assim como de uma série de adaptações dos órgãos genitais internos da mãe. A segmentação refere-se às primeiras divisões celulares do embrião. -/- • _____TRANSPORTE DO OVÓCITO -/- O ovócito que é liberado na ovulação, e que se encontra coberto pelas células do cumulus (figura 1), é capturado pela fímbria do infundíbulo ao aderir aos cílios. Esse processo é altamente eficaz, inclusive em espécies onde os ovários possuem várias formações globosas, por exemplo nas porcas, onde os ovidutos capturam entre 95 e 100% dos ovócitos que são ovulados. As contrações das camadas musculares do oviduto e o movimento intenso dos cílios da mucosa faz com que as secreções fluam em direção ao útero, transportando assim o complexo cumulus-ovócito. Esse transporte é relativamente rápido até alcançar a junção da ampola com o istmo (que é considerado o local de fertilização do espermatozoide e ovócito) a partir do qual se torna lento. Devido a fêmea estar em estro, o processo está sob controle endócrino, isto é, controle hormonal através do estradiol. Nas fêmeas domésticas, ao contrário das mulheres, o transporte ao longo do oviduto é extremamente eficiente, pois os embriões passam para o útero sem dificuldade, de modo que gestações ectópicas (tubárias ou em cavidade) são quase inexistentes. A égua, frequentemente, retém por muito tempo os ovócitos não fertilizados no oviduto, provavelmente devido a não secreção de substâncias (como a prostaglandina E) que poderiam favorecer sua passagem, como se propõe que aconteça com os embriões. A figura 2 apresenta um ovócito com suas respectivas estruturas. Figura 1: ovócitos de uma vaca madurados in vitro. Nota-se as células do cumulus que os recobre. -/- Figura 2: estruturas do ovócito de uma ovelha que se encontra na etapa de ovócito secundário, depois da ovulação. Elaborado pelo autor. • _____TRANSPORTE ESPERMÁTICO -/- Para que os espermatozoides sejam capazes de fertilizar o ovócito, deverão sofrer uma série de mudanças bioquímicas e morfológicas ao longo de sua passagem pelo aparelho reprodutor tanto masculino como feminino. Uma vez produzidos na parede do túbulo seminífero, os espermatozoides são liberados em direção ao lúmen tubular e transportados passivamente para uma estrutura ramificada conhecida como rede testicular (rete testis). Dessa rede são conduzidos em direção ao epidídimo passando através de 10 a 20 ductos eferentes localizados no polo superior do testículo. O epidídimo é dividido em três seções denominadas cabeça, corpo e cauda; é constituído por um só ducto longo e tortuoso que continua com o canal deferente. Ao final desemboca nas ampolas seminais, no ducto ejaculatório e na uretra. As funções do epidídimo são as de maduração, transporte e armazenamento dos espermatozoides. O transporte através do epidídimo é lento, aproximadamente de 10 dias em touros, e segue sendo passivo. Os espermatozoides tomados da cabeça do epidídimo são ainda imaturos e incapazes de fertilizar, enquanto os armazenados na cauda são completamente maduros. Durante o trânsito pelo epidídimo os espermatozoides adquirem motilidade e o potencial para fertilizar, o acrossomo é remodelado e a gota citoplasmática migra para o flagelo e é liberada. Quando um macho ejacula com muita frequência, é possível observar espermatozoides com gota citoplasmática no sêmen, já que não há tempo suficiente para que completem sua maduração. Os espermatozoides são expulsados fora do organismo durante a cópula, na masturbação ou em emissões espontâneas. Na ejaculação, os espermatozoides que se encontram suspensos nos fluidos do testículo e do epidídimo, misturam-se ao chegar na uretra com as secreções das glândulas acessórias para formar o sêmen. Essas secreções denominadas plasma seminal, proporcionam substâncias para manter o metabolismo energético das células espermáticas, e integram suas membranas elementos que impedem uma capacitação prematura. Durante a cópula, o sêmen é depositado na vagina ou no útero, variando entre as espécies (tabela 1). Na monta natural, geralmente, o serviço ocorre no momento propício, já que está definido pela fase do ciclo estral em que a fêmea é receptiva ao macho. No entanto, na inseminação artificial, é o macho que deverá determinar o momento ideal, e para isso é importante considerar a vida média dos gametas, ao qual é muito curta no caso dos ovócitos (tabela 1). -/- Tabela 1: local de depósito do sêmen, volume da ejaculação e vida média dos gametas. Espécie Local de deposição Volume (ml) Velocidade de ejaculação Vida média do ovócito Vida média do espermatozoide Bovinos Intravaginal 2 1 – 3 seg 8 h 30 – 48 h Caninos Intravaginal 2 – 30 (10 média) 6 – 45 min 48 – 72 h 9 – 11 d Equinos Intrauterina 50 – 200 20 – 60 seg 6 – 8 h 72 – 120 h Humanos Intravaginal 3 - - 5 – 6 d Ovinos Intravaginal 1 1 – 2 seg 16 – 24 h 30 – 48 h Suínos Intrauterina 200 – 400 5 – 20 min 8 – 10 h 24 – 48 h Onde d = dias; h = horas. A vida fértil do ovócito é muito curta, portanto, o momento do serviço é de grande importância para obter altos índices de fertilização. -/- Independentemente do local em que os espermatozoides sejam depositados no aparelho reprodutor feminino, serão expostos às secreções genitais e sofrem uma série de mudanças em seu trajeto até o local de fertilização antes de penetrar no ovócito. Nas espécies em que o sêmen é depositado na parte cranial da vagina, uma parte do mesmo penetra através da cérvix, enquanto outra parte é eliminada do aparelho genital da fêmea, em pouco tempo, através do fluxo retrógrado. O meio vaginal não é adequado e imobiliza os espermatozoides em pouco tempo, pelo qual deverão entrar no útero onde o ambiente é mais propício. O transporte espermático na fêmea é o resultado da alta contratilidade, do movimento ciliar e o fluido do aparelho genital durante o estro, ao qual está sob controle endócrino e do sistema nervoso. Esse transporte é favorecido pelas características especiais do muco estral, cujas moléculas formam uma espécie de canais que facilitam a passagem dos espermatozoides. Pelo contrário, durante a fase lútea seu transporte é dificultado. Além do meio vaginal, a cérvix também atua como barreira natural para limitar a passagem dos espermatozoides, diminuindo assim a possibilidade de polispermia. Funciona, também, como um filtro que seleciona os espermatozoides aptos dos que não o são, uma vez que somente os primeiros possuem uma motilidade vigorosa que lhes permite passar pelo muco altamente hidratado. Na vaca a cérvix é considerada como um reservatório espermático. Quando atravessam a cérvix, os espermatozoides seguem sua deslocação tanto por movimento próprio como pelas contrações uterinas e tubárias. Nas espécies em que o local de depósito do sêmen do macho na fêmea é intrauterino, como equinos, a principal barreira que os espermatozoides enfrentam é a união ou junção útero-tubárica. A união entre o útero e o oviduto (istmo do oviduto) serve como reservatório funcional dos espermatozoides nas espécies domésticas. Poucos minutos depois da cópula, é possível encontrar alguns espermatozoides no oviduto, que é conhecido como fase de transporte rápido. No entanto, esses espermato-zoides não são os que participam do processo de fertilização e podem apresentar certas anormalidades. Existe um segundo tipo de transporte denominado fase sustentada, a qual consiste na migração prolongada dos espermatozoides em direção as partes mais craniais do aparelho genital feminino, que conduzem à colonização do reservatório funcional e na liberação gradual dos espermatozoides dos reservatórios espermáticos, incluindo este último. O reservatório do istmo fornece aos espermatozoides um ambiente propício, protegendo-os contra a fagocitose, prolongando assim a sua viabilidade. Aqui os esper-matozoides permanecem aderidos a superfície das células ciliadas do epitélio até a finalização de sua capacitação, depois da qual são liberados quando alteram seu padrão de motilidade flagelada, fenômeno conhecido como hipermotilidade, e migram em ondas em direção ao local da fertilização. Dos milhões de espermatozoides ejaculados, somente alguns milhares alcançarão o istmo do oviduto e um número sumamente pequeno será encontrado nas imediações do ovócito no momento da fertilização. -/- • _____CAPACITAÇÃO ESPERMÁTICA E REAÇÃO ACROSSÔMICA -/- A capacitação é um processo gradual e essencial para a fertilização. Os espermato-zoides devem passar por um dado tempo de “incubação” no aparelho genital da fêmea e sofrer uma série de mudanças antes de serem capazes de fecundar o ovócito. A capacita-ção dos espermatozoides começa quando eles entram em contato com as secreções do aparelho genital da fêmea e termina no istmo do oviduto. Durante esse percurso a superfície da cabeça do espermatozoide se modifica já que algumas moléculas como o colesterol são removidas da membrana plasmática, aumentando sua fluidez e alterando suas propriedades bioquímicas. Entre outras coisas, essa reorganização dos lipídios facilita a entrada de cálcio extracelular pelos canais iônicos e ocasiona a desestabilização da membrana, tornando-a mais fusogênica. Também são eliminados outros fatores (conhecidos genericamente como fatores descapacitantes) que expõem receptores membranais indispensáveis para a realização da união entre o espermatozoide e o ovócito durante a fertilização. Os espermatozoides capacitados apresentam um padrão de hipermotilidade e uma maior atividade metabólica, características que devem obter para que sejam capazes de penetrar as camadas do ovócito. A capacitação é necessária para que ocorra a reação acrossômica. A reação acrossomal ou acrossômica (RA) é um fenômeno de exocitose que é desencadeada pela ligação entre as proteínas e receptores localizados na membrana do espermatozoide e a zona pelúcida do ovócito (figura 3). Envolve a fusão da membrana plasmática do espermatozoide com a membrana externa de seu acrossomo; formando, assim, pequenas vesículas, cujo conteúdo de enzimas hidrolíticas, como a hialuronidase e a acrosina, é liberado em direção ao seu exterior, facilitando a penetração da zona pelúcida. A adesão do espermatozoide à zona pelúcida é específica da espécie e depende de glicoproteínas presentes na membrana dos ovócitos maduros; a cabeça do espermatozoide se une com essas glicoproteínas mediante receptores específicos e atravessam em direção oblíqua a zona pelúcida até chegar ao espaço perivitelino. Nos mamíferos, particularmente em roedores e nos humanos, algumas dessas glicoproteínas são conhecidas como ZP1, ZP2 e ZP3. Pensa-se que a união da membrana espermática a esta última é o que desencadeia a reação acrossomal. Posteriormente, as vilosidades do ovócito entram em contato com o espermatozoide e a membrana presente na secreção equatorial da cabeça do espermatozoide, que tem proteínas fusogênicas específicas, une-se a membrana plasmática do ovócito, fusionando-se e permitindo a entrada do núcleo espermático ao citoplasma (figura 4). A capacitação dos espermatozoides é fundamental para o êxito da fertilização, uma vez que apenas os aptos estarão perfeitos e íntegros para que a união e percepção das glicoproteínas presentes nos mesmos possam ocorrer de forma ideal para a penetração deste no ovócito, fertilizá-lo e formar um zigoto. Figura 3: estrutura do espermatozoide. Nota: a membrana é acrossomal e não acrossonal. Elaborado pelo autor. -/- Figura 4: eventos que ocorrem durante a fertilização. A – depois de passar através das células do cumulus, o espermatozoide entra em contato com a zona pelúcida, onde receptores da membrana plasmática reconhecem as proteínas da zona pelúcida; B – é desencadeada, então, a reação acrossomal; C – para que o espermatozoide penetre na zona pelúcida; D – ao atravessar a zona pelúcida e entrar no espaço perivitelino, a cabeça do espermatozoide entra em contato com a membrana vitelina; ambas membranas se fusionam graças ao reconhecimento de proteínas fusogênicas que estão no segmento equatorial. E – ocorre, então, que o núcleo do espermatozoide penetra no citoplasma do ovócito; uma das consequências da fusão da membrana com a cabeça do espermatozoide é a reação cortical, em que os grânulos corticais do ovócito liberam seu conteúdo em direção ao espaço perivitelino, o que resulta na alteração da estrutura da zona pelúcida e da membrana vitelina para bloquear a polispermia. Elaborado pelo autor. • _____CONSEQUÊNCIAS DA PENETRAÇÃO -/- Bloqueio da polispermia -/- A fusão das membranas dos gametas durante a penetração permite a entrada da fosfolipase C zeta (PLCζ) – fator solúvel que provém do espermatozoide – que desencadeia liberação e oscilações de Ca2+ no retículo endoplasmático. Isso, por sua vez, provoca a migração e fusão dos grânulos corticais, com a consequente liberação de enzimas, que mudam tanto a estrutura da zona pelúcida (por exemplo a inativação de ZP3), como a da membrana vitelina. Desse modo, impede-se que outros espermatozoides as penetrem, evitando assim a polispermia (figura 5). -/- Figura 5: os grânulos corticais que encontram-se na periferia do citoplasma, debaixo da superfície da membrana do ovócito, migram e fusionam-se sobre a mesma, liberando seu conteúdo no espaço perivitelino imediatamente depois da penetração do espermatozoide. A – como conse-quência, tanto a membrana vitelina como a zona pelúcida são modificadas, impedindo a entrada de mais espermatozoides, o que evita a polispermia. B – o ovócito retoma, então, sua segunda divisão meiótica, que ocasiona a expulsão do segundo corpúsculo polar, e na formação dos pronúcleos feminino e masculino. C – continuando, os pronúcleos migram, suas membranas se dissolvem e os cromossomos de ambos se condensam e se unem. Completa-se, assim, a singamia e forma-se, então, o zigoto. Elaborado pelo autor baseado nos livros de embriologia e fisiologia da reprodução conforme vida bibliografia. -/- Ativação do ovócito e formação de pronúcleos -/- Na maioria das fêmeas domésticas, com exceção da cadela, o ovócito encontra-se suspenso na metáfase II da segunda meiose no momento da ovulação. A entrada da PLCζ do espermatozoide no citoplasma do ovócito, e a consequente liberação de Ca2+, faz com que o ovócito se ative, terminando a segunda divisão meiótica e expulse o segundo corpo polar. Posteriormente, o material nuclear do ovócito se reorganiza para formar o pronú-cleo feminino. Entretanto, a membrana nuclear do espermatozoide se dissolve, a cromatina descondensa-se, as protaminas são substituídas por histonas, e se forma uma nova membrana nuclear, dando lugar ao pronúcleo masculino. -/- Singamia -/- Uma vez que os pronúcleos feminino e masculino são formados, migram para o centro do ovócito, aproximam-se, suas membranas se dispersam e os cromossomos paternos e maternos se associam, com o qual recupera a condição diploide e dá origem ao zigoto (figura 5). -/- • _____DESENVOLVIMENTO EMBRIONÁRIO -/- Poucas horas após a fertilização ocorre a primeira divisão do zigoto em duas, depois em 4, 8, 16 e 32 células, denominadas blastômeros. Essas divisões mitóticas são conhecidas como divisões de segmentação ou clivagem (figura 6), uma vez que são realizadas sem aumento do citoplasma, de modo que com cada divisão os blastômeros se tornam menores. A partir das 16 células o embrião se chama mórula, e é visto como uma massa celular compacta. A compactação deve-se a informação de proteínas de ligação entre os blastômeros. O embrião acumula líquido em seu interior, formando uma cavidade denominada blastocele. Esse processo é conhecido como blastulação; o embrião, assim, passa a ser chamado de blastocisto. Nessa etapa é possível diferenciar duas populações de células embrionárias: a massa celular interna, embrioblasto ou botão embrionário (que dará origem ao embrião), e a massa celular externa, células superficiais ou trofoblasto, do qual se originam a maioria das membranas fetais. Ao continuar a multiplicação das células e a acumulação de líquido, o blastocisto aumenta de tamanho, convertendo-se em blastocisto expandido. A zona pelúcida torna-se mais fina e, finalmente, o embrião eclode; isto é, o embrião é liberado da zona pelúcida. -/- • _____FERTILIZAÇÕES ATÍPICAS -/- Polispermia -/- É a penetração de dois ou mais espermatozoides no óvulo. Esta condição é letal nos mamíferos, já que o número cromossômico desse zigoto é maior que 2n, o envelheci-mento do ovócito da porca, como consequência do serviço tardio, favorece a apresentação da polispermia, por isso, é comum ver números de nascimentos de média de 10 a 12 leitões por parição nessa espécie. Nas aves, no entanto, a penetração de mais de um espermatozoide é normal, embora apenas um pronúcleo masculino se formará, fundindo-se com o feminino. -/- Figura 6: etapas do processo de segmentação/clivagem. Fonte: aula de reprodução da professora Domenica Palomaris, UFT. -/- Ginogênese -/- É o desenvolvimento de um embrião a partir de um óvulo normal fecundado por um espermatozoide, mas sem a fusão dos cromossomos masculinos com os da fêmea. A função do espermatozoide, nesse caso, é a de ativação do ovócito para que este inicie seu desenvolvimento, mas não há fusão com o núcleo do espermatozoide. Ocorre em plantas, em nematódeos e em algumas espécies de peixes, por exemplo a Poecilia formosa, espécie em que os ovócitos das fêmeas são ativados por machos de outra espécie relacio-nada. -/- Partenogênese -/- Consiste no desenvolvimento do embrião sem a participação do espermatozoide. Ocorre em alguns insetos, o zangão, por exemplo, é partenogenético. Também pode apresentar-se em perus, cujos embriões são machos e, geralmente, morrem antes da eclosão. -/- • _____GÊMEOS -/- Existem dois tipos de gêmeos: idênticos ou monozigóticos e os não idênticos ou dizigóticos. -/- Idênticos ou monozigóticos -/- Originam-se do mesmo zigoto, sendo assim, possuem o mesmo genótipo e um fenótipo similar e, portanto, são do mesmo sexo. Em laboratório é possível gerá-los ao seccionar uma mórula em duas ou mais partes, por meio de um micromanipulador. Uma vez que possuem o mesmo genótipo, os produtos resultantes são clones. -/- Não idênticos ou dizigóticos -/- Provêm da fertilização de dois óvulos distintos por espermatozoides diferentes. Possuem, portanto, genótipos e fenótipos diferentes, e podem ser de sexo diferente. -/- REFERÊNCIAS BIBLIOGRÁFICAS -/- ALBERTS, Bruce et al. Biologia molecular da célula. Artmed Editora, 2010. AUSTIN, Colin Russell; SHORT, Roger Valentine. Reproduction in Mammals, Book I: Germ Cells and Fertilization. 1982. BEARDEN, Henry Joe et al. Reproducción animal aplicada. México: Manual Moderno, 1982. BOSCH, P.; WRIGHT JR, R. W. The oviductal sperm reservoir in domestic mammals. Archivos de medicina veterinaria, v. 37, n. 2, p. 95-105, 2005. CAPALLEJAS, Roberto Brito; RODRÍGUEZ, Lourdes Tagle. Fisiología de la reproducción animal: con elementos de biotecnología. Editorial Félix Varela, 2010. CROXATTO, Horacio B. Physiology of gamete and embryo transport through the fallopian tube. Reproductive biomedicine online, v. 4, n. 2, p. 160-169, 2002. CUPPS, Perry T. (Ed.). Reproduction in domestic animals. Elsevier, 1991. DIEDRICH, Smidt et al. Endocrinología y Fisiología de la Reproducción de los Animales Zootécnicos. 1972. DUKES, Henry Hugh; SWENSON, Melvin J.; REECE, William O. Dukes fisiologia dos animais domésticos. Editora Guanabara Koogan, 1996. FERREIRA, A. de M. Reprodução da fêmea bovina: fisiologia aplicada e problemas mais comuns (causas e tratamentos). Juiz de Fora: Minas Gerais–Brasil, p. 422, 2010. HAFEZ, Elsayed Saad Eldin; HAFEZ, Bahaa. Reprodução animal. São Paulo: Manole, 2004. HUNTER, Ronald Henry Fraser. The Fallopian tubes in domestic mammals: how vital is their physiological activity?. Reproduction Nutrition Development, v. 45, n. 3, p. 281-290, 2005. HUNTER, R. H. F.; RODRIGUEZ‐MARTINEZ, H. Capacitation of mammalian spermatozoa in vivo, with a specific focus on events in the Fallopian tubes. Molecular Reproduction and Development: Incorporating Gamete Research, v. 67, n. 2, p. 243-250, 2004. GALINA, Carlos; VALENCIA, Javier. Reproducción de los animales domésticos. 2006. HOPPER, Richard M. (Ed.). Bovine reproduction. John Wiley & Sons, 2014. HYTTEL, Poul; SINOWATZ, Fred; VEJLSTED, Morten. Embriologia veterinária. Elsevier Brasil, 2012. KÖLLE, Sabine; REESE, Sven; KUMMER, Wolfgang. New aspects of gamete transport, fertilization, and embryonic development in the oviduct gained by means of live cell imaging. Theriogenology, v. 73, n. 6, p. 786-795, 2010. MCKINNON, Angus O. et al. (Ed.). Equine reproduction. John Wiley & Sons, 2011. OLIVERA, Martha et al. El espermatozoide, desde la eyaculación hasta la fertilización. Revista Colombiana de Ciencias Pecuarias, v. 19, n. 4, p. 426-436, 2006. PLANT, Tony M.; ZELEZNIK, Anthony J. (Ed.). Knobil and Neill's physiology of reproduction. New York: Academic Press, 2014. RODRIGUEZ-MARTINEZ, Heriberto. Role of the oviduct in sperm capacitation. Theriogenology, v. 68, p. S138-S146, 2007. SAAVEDRA LEOS, María Dolores. Estudio de la composición de los gránulos corticales y del oolema de ovocitos porcinos y bovinos madurados y fecundados in vitro. Proyecto de investigación:, 2010. SENGER, Phillip L. et al. Pathways to pregnancy and parturition. Current Conceptions, Inc, 2004. SUAREZ, Susan S. The oviductal sperm reservoir in mammals: mechanisms of formation. Biology of Reproduction, v. 58, n. 5, p. 1105-1107, 1998. VALENCIA MÉNDEZ, Javier de J. Fisiología de la reproducción porcina. 1986. -/- FIXAÇÃO DO ASSUNTO -/- 1. Defina e diferencie fertilização e segmentação. -/- 2. Disserte sobre o transporte do ovócito. Por que no istmo o transporte do ovócito se torna lento? -/- 3. Fale sobre a eficiência do transporte do gameta no trato genital da fêmea e sobre a retenção de ovócitos pela égua. -/- 4. Defina as células do cumulus e explique seu papel sobre o ovócito. -/- 5. Quais as estruturas que compõem o ovócito? Qual a função de cada uma? -/- 6. Todos os ovócitos das fêmeas domésticas são iguais? Justifique. -/- 7. Disserte sobre o papel dos hormônios e das secreções sobre o transporte do ovócito no trato genital da fêmea. -/- 8. Disserte sobre o transporte do espermatozoide. Por que há transporte rápido e lento dos espermatozoides? -/- 9. Fale sobre o local de deposição do sêmen nas espécies domésticas e como esse influen-cia na eficiência da reprodução. -/- 10. Por que a vida média do espermatozoide canino é maior que as outras espécies? -/- 11. Quais as mudanças que os espermatozoides precisam sofrer para conseguir êxito na fertilização e por quê? -/- 12. Defina fase de transporte rápido, fase sustentada e hipermotilidade. Qual a importân-cia de cada um? -/- 13. Como ocorre a reação acrossômica? -/- 14. O que é polispermia e como ocorre seu bloqueio? 15. Como ocorre a ativação do ovócito e como se formam os pronúcleos? -/- 16. O que é e qual a importância da singamia? -/- 17. Disserte sobre as divisões de segmentação e qual a importância destas para o êxito da reprodução. -/- 18. Defina e diferencia mórula e blastocisto. -/- 19. Defina e diferencie ginogênese e partenogênese. -/- 20. Qual o papel endócrino e quais hormônios e enzimas atuam sobre o transporte de gametas, sobre a fertilização e sobre a segmentação? (shrink)

Language is nothing but human subjects in as much as they speak, say and know. Language is something coming from the inside of the speaking subject manifest in the intentional meaningful purpose of the individual speaker. A language, on the contrary, is something coming from the outside, from the speech community, something offered to the speaking subject from the tradition in the technique of speaking. The speech act is the performance of an intuition by the subject, both individual and social.

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)

From the second half of the last century, there has been a widespread view in the Anglophone world that Kant’s transcendental deduction (aka TD) aims to vindicate our common-sense view of the world as composed of public and objective particulars against some unqualified forms of skepticism. This widespread assumption has raised serious doubt not only about the success of TD but also about the very nature of its argument in both editions of the Critique. Yet, if there is a connection (...) between TD and global skepticism, the intriguing question is: Who is this skeptic? According to Strawson, “the skeptic” is a hypothesis of a purely sense-datum experience. In contrast, the fact that TD turns on the key notion of self-consciousness has induced several other scholars to assume that the skeptic is no none but a Cartesian external-world skeptic. None of those readings find textual support or are compatible with the very structure of the first Critique. The question is: Does this mean that TD aims only to undermine empiricism, as Guyer suggests? I do not believe so. I am pretty convinced that TD addresses a peculiar form of global skepticism, namely “Hume’s challenge to reason.” Assuming that we cognize (erkennen) and experience (erfahren) appearances as objects as a requirement of Newtonian physics, TD aims to provide a justification of the principle that nature is uniform that is superior to Hume’s justification. (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)

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)

This brief paper is devoted to criticizing the widespread reading of Kant’s first Critique, according to which reference to subject-independent objects is “constituted” by higher-order cognitive abilities (concepts). Let us call this the “constitutional view.” In this paper, I argue that the constitutional reading confuses the un-Kantian problem of how we come to represent objects (which I call the intentionality thesis), with the quite different problem of how we cognize (erkennen) (which I call the “cognition thesis”) that we do represent (...) objects, that is, things that exist independently of the subject. (shrink)

This article presents a dual purpose: to carefully consider objections against the existentialist conception of the content of visual experience and to develop and defend a version of it that avoids such objections, specifically addressing the so-called "problem of particularity." The main thesis is that the existential content of visual experience should be understood as relativized, being incomplete content (rather than classical, complete propositions), modeled as a function of the sextuple of the object, agent, time, place, causal relation, and world (...) to absolute values. (shrink)

In this paper, I argue against the phenomenal concept strategy (henceforth PCS) and in favor of what Chalmers has called type-A materialism ([2006], [2010] p. 111). On her release, Mary makes no cognitive discovery at all, not even a thin, non-possibility-eliminating discovery, as Tye has recently claimed [2012]. When she is imprisoned, Mary already knows everything that is to be known about the phenomenal character of her experiences. What Mary acquires is a new non-cognitive and nonconceptual representation.

This paper is a new non-descriptivist defense of non- conceptualism, based on a new interpretation of Kant’s metaphysics of concepts. We advance the following claim: What distinguishes non-conceptual from conceptual singular representations is the way partial representations of the object’s features are integrated into the whole representation of the object: while at the non-conceptual level this integration takes the form of images of the object’s features that are stored and projected, at the conceptual level this integration takes the form of (...) the recognition that those features are properties of a same object, what Kant calls “synthetic unity of Aperception”. (shrink)

Content nonconceptualism and State conceptualism are motivated by different readings of what I want to call here Bermúdez’s conditions on content-attribution (2007). In one read- ing, what is required is a neo-Fregean content to solve problems of cognitive significance at the nonconceptual level (Toribio, 2008; Duhau, 2011). In the other reading, what is required is a neo-Russellian or possible-world content to account for how conspecifics join attention and cooperate, contemplating the same things from different perspectives in the same perceptual field. (...) The solution to this apparent contradiction is the rejection of the real content view and the adherence to what I call here Content-pragmatism: there is no such thing as the content of experience. According to content pragmatism, “proposition” is not as real as a mental state, but rather, it is a term of art that semanticists use, as a matter of theoretical convenience, to classify mental states. What follows from Content-pragmatism is Content- pluralism: there are so many contents that are required to meet Bermúdez’s condition on content attribution. Because both criticism and the defense of State conceptualism overlook the real scope of Bermúdez’s condition on content attribution, they are ineffective. In this paper, I will argue that the opposition between State and Content nonconceptualism is a real one, but only a relative one, that is, relative to the opposite constraints to be met. If we want to solve the problems of cognitive significance, the best we can do is to let the content of experience be modeled as neo-Fregean content, namely, a compound of nonconceptual modes of presentation of objects and properties. In this case, Content-nonconceptualism prevails. In contrast, if we want to account for how conspecifics join attention to the same entities in the same perceptual field from different perspectives, the best we can do is let the content of experience be modeled either as a structured Russellian proposition or as a function from possible worlds to truth-values. In this case, State-nonconceptualism prevails. (shrink)

Formal ontology as it is presented in Husserl`s Third Logical Investigation can be interpreted as a fundamental tool to describe objects in a formal sense. It is presented one of the main sources: chapter five of Carl Stumpf`s Ûber den psycholoogischen Ursprung der Raumovorstellung (1873), and then it is described how Husserlian Formal Ontology is applied in Fifth Logical Investigation. Finally, it is applied to dramatic structures, in the spirit of Roman Ingarden.

This essay aims to expose the metaphysical underpinnings of enactivism. While enactivism relies heavily on rejecting the traditional mind-body problem by excluding the familiar thought experiments that favor phenomenal dualism, the crucial point that is overlooked is instead the brain-body problem, specifically the crucial interaction between the brain and the bodily limbs in their embodied activities of perception and cognition. If enactivism is correct, differences in sensory experience necessarily entail differences in embodied activity—this is the metaphysical core of enactivism, which (...) we think is entirely wrong. We will argue that a physical or biological body ("Körper") is configured or shaped into a perceptually living body ("Leib") by brain patterns and not by embodied activity or by the so-called sensorimotor contingencies. Undoubtedly, embodied actions influence sensory perception. However, since variations in embodied actions do not necessarily lead to variations in sensory experience, they are not metaphysically constitutive of these experiences (supervenience claim). (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)

Kant was never satisfied with the version of his “Refu- tation” published in 1787 (KrV, B 275-279). His dissatisfaction is already evident in the footnote added to the preface of the second edition of the Critique in 1787. As a matter of fact, Kant continued to rework his argument for at least six years after 1787. The main exegetical problem is to figure out who is the target of the “Refutation”: a non-skeptic idealist, a global skeptic of Cartesian provenance or (...) both. In this last case, the related problem is to know whether either of them is the Cartesian skeptic of the first Meditation, the idealist skeptic of the second Meditation and first part of the third Meditation, or some other non-skeptic idealist. I present and defend a new reconstruction of Kant’s “Refu- tation” as a successful argument against Mendelssohnian idealism of Cartesian provenance. This defense is based on a simple logical sketch of the proof provided by Dicker but essentially modified in light of Dyck’s insight about Kant’s opponent. How shall I support my reading? First, by appealing to overwhelming textual evidence according to which the proof is of the existence of mind-independent things, showing that Kant’s main opponent is Mendelssohn’s idealism of Cartesian provenance. Finally, I support my reading by showing that Kant’s “Refutation” is doomed to fail against all forms of global skepticism but is quite successful against Mendelssohn’s idealism. (shrink)

This paper is a new defense of type-B materialism against Jackson’s knowledge argument (1982) inspired by the Kantian main opposition between concepts and sensible intuitions. Like all materialists of type B, I argue that on her release from her black-and-white room, Mary makes cognitive progress. However, contrary to the so-called phenomenal concept strategy (henceforth PCS), I do not think that such progress can be accounted for in terms of the acquisition of new concepts. I also reject Tye’s recent account of (...) Mary’s cognitive progress as the acquisition of a “thing-knowledge.” What is crucial is not the Russellian opposition between knowing things and knowing truths but rather the Kantian opposition between conceptual and nonconceptual representations of the same thing (property). Mary’s phenomenal knowledge is here accounted for as the result of the cooperation of her newly acquired nonconceptual representation of the same phenomenal redness (the same thing) she had a conceptual representation before. As that new nonconceptual representation carries information about the same physical property she already represented but is now coded in analog rather than in digital form (Dretske, 1981), that nonconceptual representation can be accounted for in physical terms as Chalmers’s Master argument requires. Nevertheless, that representation can account for Mary’s cognitive progress since it carries information coded in analog form, something that the imprisoned she could never possess. (shrink)

My account of the causal role of consciousness in a physical world is modeled on Dretske’s celebrated explanation of the causal role of beliefs (something that Dretske himself never offered). First, behavior must be understood as a (broadly individuated) process that begins with some external stimulus causing some neurological event C, and ends with causing a bodily movement M (e.g., the Kennedy assassination is a process that begins with Oswald pulling the trigger at 12:30pm CST on November 23 in 1963 (...) in Dallas, Texas, but only ends half an hour later when Kennedy is pronounced dead at 1 pm CST). The internal neurological event C causes bodily movement M, but only by virtue of being recruited by natural selection to represent the instantiation of some external property F when properly stimulated under normal circumstances. But the reason why C causes M lies in the fact that C represents the instantiation of the external property F. I withdraw my hand from a hot surface because the activation of nociceptive specific neurons in my parietal lobe (together with the activation of neuronal patterns in my motor cortices) was recruited by natural selection to represent the tissue damage in my hand. The activation of nociceptive specific neurons causes my hand to withdraw but for the reason that it represents tissue data at the time that I felt pain in my hand. (shrink)

This paper is a defense of the so-called phenomenal-concept strategy, based on a new view of phenomenal concepts as special de re modes of presentation of the phenomenal character of experience. Phenomenal concepts can be explained in physical terms as mental particulars (as phenomenal files) created in the individual’s mind to pick out the phenomenal character of experience by representing certain physical properties as those represented by the experiences themselves (metarepresentation). They are individuated by two fundamental relations: the perceptual (acquaintance) (...) relation the creature bears to the physical properties represented by its own experience and the relation the creature bears to itself (self-acquaintance) as the subject undergoing those experiences. Mary’s newly acquired phenomenal file presents the phenomenal character of her new experience of red by means of the new relation she bears to herself as the creature standing in the experiential relation to the color red. (shrink)

This paper is both systematic and historical in nature. From a historical viewpoint, I aim to show that to establish Wittgenstein’s claim that “an ‘inner process’ stands in need of outward criteria” (PI §580) there is an enthymeme in Wittgenstein’s private language argument (henceforth PLA) overlooked in the literature, namely Wittgenstein’s suggestion that both perceptual and bodily experiences are transparent in the relevant sense that one cannot point to a mental state and wonder “What is that?” From a systematic viewpoint, (...) I aim to show that Wittgenstein’s PLA teaches us that the prevailing picture of the nature of phenomenal concepts (henceforth PCs) is upside down: we can only introspectively know what is going on inside our heads after we learn of what is going on outside (PI §580). In this regard, I aim to defend two associate claims against the prevailing view of PCs on the basis of PLA. First, by means of transparency, I aim to show that there is no de re awareness of our private sensation that could determine the meaning of sensation words; for example, I am never aware of the phenomenal blueness of my experience of something blue. The second associated claim is that introspective self-knowledge of our private sensations is always de dicto. We can only know introspectively that phenomenal blueness is the phenomenal character of the experience we are undergoing after we have learned that (de dicto knowledge) blue is the color that usually causes us that kind of experience. Likewise, we can only introspectively know that pain is the phenomenal character of the experience we are undergoing after we have learned that pain is what usually causes some typical pain behavior. (shrink)

This paper has two aims. First, it aims to provide an adverbial account of the idea of intransitive self-consciousness and second, it aims to argue in favor of this account. These aims both require a new framework that emerges from a critical review of Perry’s famous notion of the “unarticulated constituents” of propositional content (1986). First, I aim to show that the idea of intransitive self-consciousness can be phenomenologically described in an analogy with the adverbial theory of perception. In an (...) adverbial theory of perception, we do not see a blue sense-data, but we see something blue-ly, whereas in intransitive self-consciousness, we are not conscious of ourselves when we undergo a conscious experience—instead, we experience something self-consciously. But what does this mean precisely? First, I take intransitive self-consciousness to be the first-person operator that prefixes the content of any experience that the subject undergoes, regardless of whether or not the subject is self-referred. Further, I argue that this first-person adverbial way of entertaining the content of any experience in Perry’s revised framework fixes the subject as part of the circumstance of the evaluation of the content of her own experience. We can only evaluate whether the content is veridical or falsidical relative to the subject undergoing the experience. This is referred to here as “self-concernment without self-reference.” When I am absorbed in reading a book, I do not self-represent my own experience of reading a book, let alone see myself as a constituent of the content of this experience. Even so, I experience that reading self-consciously in the precise sense that I do belong to the circumstance of the evaluation of the selfless content of my experience of reading the book. The content of the experience of reading a book is simply a propositional function, true or false of myself. (shrink)

In this paper, I intend to show that it’s a serious mistake to construe the role of sensible representation in Kant’s work as a nonconceptual content (in the contemporary and technical sense of “content”), which, like a mental indexical would refer to what appears in space and time in the so-called de re form. The interpretation I advance and further support is this: without possessing a representational content, sensible representation must be understood as the basic epistemic relation between the subject (...) and the physical entities in space and time she is acquainted with. Immediate reference of sensible intuition to their respective objects is to be understood as a form of knowledge by acquaintance. I call my own interpretation relationalism. (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.

In this critical review of Robert Hanna's ingenious book (2006), I aim to support Hanna‟s main insightful reading of Kant, namely what he calls “a priori truth with a human face," without appealing to Kant's divide between a priori and a posteriori and analytic and synthetic truths. My suggestion is that transcendental propositions are necessary neither in the usual epistemological sense that analytic propositions are, let alone in the metaphysical sense that some empirical propositions are. Instead, they are necessary in (...) the theoretical domain in the weak alternative sense that they make possible the empirical recognition of appearances as an object as an indispensable condition of our self-consciousness experience, and they are a priori in the practical domain in the sense that their truth is vital for our self-comprehension as human beings. (shrink)

Este artículo pretende ofrecer una visión alternativa tanto de la lectura conceptualista tradicional de Kant como de la nueva lectura no conceptualista. En contra de las lecturas conceptualistas tradicionales sostengo que confunden las condiciones para la representación sensible de los objetos (tesis de la intencionalidad) con las condiciones para el reconocimiento (Erkenntnis) de que representamos objetos mediante intuiciones sensibles (tesis del reconocimiento). En contra de las lecturas no conceptualistas sostengo que no distinguen el no conceptualismo -propio de la filosofía contemporánea (...) de la mente- de las tesis antiintelectualistas de Kant. Tales lecturas fracasan porque se equivocan al atribuir un contenido representacional a las intuiciones sensibles de Kant. Las intuiciones sensibles, según Kant, no presentan el mundo según ciertas condiciones de satisfacción que podrían o no ser satisfechas por el mundo. Por lo tanto, no son ni verídicas ni non verídicas. Se limitan a presentar el mundo de una manera directa, independiente de los conceptos, lo que llamaré aquí la presentación de re de los objetos. En mi lectura, la oposición entre intuiciones y conceptos es la oposición entre la presentación de re de um objeto y el reconocimento proposicional de dicto de que representamos algo como um objeto. (shrink)

I aim to develop an alternative theoretical model for the direct cognition of the elementary states of others called the theory of interaction (henceforth TI), also known as the “second person” approach. The model I propose emerges from a critical reformulation of the displaced perception model proposed by FRED DRETSKE (1995) for the introspective knowledge of our own mental states. Moreover, against Dretske, I argue that no meta-representation (second-order representation of a first-order representation as a representation) is involved in the (...) process, but only concepts, rudimentary enough, of others’ emotions: contentment and discontentment. Strictly speaking, all the empirical data and findings in developmental psychology only support the hypothesis that children in the pre-linguistic phase understand perceptual facial expressions of contentment and discontentment. My proposal consists of the following thesis: The direct perception of others' elementary mental states is not a perception in the usual sense of the word according to which we just perceive things, but rather a “perception of facts.” But how to understand that? It is nothing more than an epistemological process that is automatic and reliable at the same time. It is reliable in the epistemological-externalist sense of not involving any inference, reflection, theorizing or epistemic justification: a baby’s perception of her mom’s facial expression of discontentment (in the usual sense of “per- ception”) does not constitute a reason supporting the belief that mommy is discontent. The very word “belief” should not be understood in the usual sense but rather in the sense of a natural inclination for truth. But at the same time, it is automatic because the belief is a result or output of a process entirely subliminal and computational in nature. Thus, for instance, a child perceives that her mom is discontent (fact-awareness) by perceiving consciously (“perception” in the usual sense) a facial expression of discontentment (sensorial input) but recognizes that her mother is discontent as a result (output) of an entirely subliminal process. (shrink)

I argue that my non-dual epistemic-phenomenalist view is the one that best harmonises my interpretation of the Fourth Paralogism with the widely shared reading of the Refutation of Idealism that I sketched and defended above. The bottom line of my view is a clear distinction between the metaphysical and epistemological sides of Kantian idealism. Again, according to my non-dual-epistemic-phenomenalism, the mundus sensibilis and mundus intelligibilis are epistemologically distinct ways of considering the metaphysically identical outside world. Appearances are nothing but the (...) way the things in themselves appear or exist inside our sensible minds as mere representations. In this sense, I reject both the two-world view, the two-aspect view, and Allais’s deflationary anti- phenomenalist reading. (shrink)

How does one know the phenomenal character of one’s own experience? I aim to present and defend a new view of the epistemology of qualia that addresses this issue. My view results from a reworking of Dretske’s displaced perception model. The guiding line is the key Wittgensteinian insight of his Private Language Argument, namely the claim that no inner perception of qualia can justify our corresponding qualia beliefs. My reworking of the original model starts with the rejection of Dretske’s representationalism, (...) as well as any other metaphysical views about the ultimate nature of qualia. The reworking of the displaced perception model I am proposing is supposed to be compatible with any available metaphysical view on qualia. The second step consists of rejecting the epistemological misgivings of the original model, namely the view that knowing qualia requires “connecting beliefs” along with “meta-representations.” Yet, Dretske’s crucial mistake has been to force knowledge of qualia into the “belief-box.” As a matter of fact, there is only one belief involved in introspecting qualia, namely the qualia-belief itself. The third final step is this. Assuming the key Wittgensteinian insight, I argue that introspection of qualia is a process that is both epis- temically reliable and automatic. It is reliable because it dispenses with justification of any beliefs whatsoever. Moreover, it is automatic because it also dispenses with reflexive connecting beliefs. For example, it takes nonconceptual awareness of the color blue as an input and automatically yields the belief that phenomenal blue is the conscious character of my experience of the sky as an output just by employing the concept BLUE. (shrink)

The purpose of this article is to offer a new view of the key relation between the content and the conscious character of visual experience. The author aims to support the following claims. First, the author rejects the qualia realist claim that conscious character is an intrinsic, nonrepresentational property of visual experience, for example, a pattern of activation of neurons. However, the author also rejects the rival widespread representationalist claim that the conscious character of visual experience is identical to, or (...) supervenes on, any specific property represented by visual experience. The positive proposal is the following. Conscious character is identical to those patterns of activation of neurons that are referentially or representationally alive. Conscious redness, for example, is a pattern of activations of neurons that is normally created only when brains of physical duplicates come in visual contact with some distal property. This pattern of activation is recruited by natural selection to represent that property. This is called meaning representationalism. (shrink)

This paper presents a new cognitivist account for the old puzzle of self-consciousness or knowing self-reference. Knowing self-reference does not rely on reflection on some putative pre-existent pre-reflexive self-consciousness, nor is it the result of a process of identification of oneself as the employer of the relevant token of “I” according to the token-reflexive rule of the first-person pronoun. Rather, it relies on the architecture of the cognitive system. By exploiting the acquaintance relation that every brain has to one’s own (...) body, a “mental file” is opened in a subliminal way to house information about oneself gained through the appropriate proprioceptors. Knowing self-reference based on self-files dispenses with self-identification because the opening of the file in the same brain/body is not a deed of the cognitive system but rather something that takes place sub-personally. (shrink)

The aim of this paper has been to present a new reconstruction of Kant’s Refutation of Idealism. I have considered several different targets of the Refutation, five of them mentioned by Kant himself. I believe that I have shown that the Refutation of Idealism is best considered only as a sound argument against Mendelssohnian subjectivist idealism, against Mendelssohnian immaterialism, and against Mendelssohnian realist idealism. First, Kant’s Refutation is a sound argument in favor of the claim that the outer things represented (...) in our minds are real rather than ideal; that is, they exist mind-independently as noumena. And second, Kant’s refutation is a sound argument for a fundamental ontology of noumena: the ultimate nature of reality and of our minds is neither material nor mental but made up of things in themselves. (shrink)

The claim that ―concepts serve as rules for the synthesis of representations‖ is understood by the mainstream of Kant‘s scholarship as if categories and concepts, in general, are conditions for the constitution of objects out of the manifold of sensations devoid of reference. That is the claim that I wish to question here. The claim comes in different flavors and formulations. Still, none of them are relevant here. I aim to provide an alternative account for the claim that ―the representation (...) makes the object possible.‖ I argue that the traditional view represents a solution to what I call the pseudo-problem of intentionality, namely to account for how a representation of something emerges out of a manifold of sensations devoid of reference. Yet, the intentionality problem is a misunderstanding of what I call the recognition thesis: the role of concepts is to provide the rules for the (mind-dependent) recognition (Erkenntnis) of the mind-independent existence of transcendental objects (noumena). ―The representation makes the object possible in the relevant sense that only by means of concepts can we recognize that what is sense-dependently represented actually exists mind-independently as transcendental objects or noumena in the negative sense. My view is embedded in the framework of what today is called ―Kantian Nonceptualism‖. Still, my view is neither a simple re-statement of Kantian Nonconceptualism nor depends on it. It is independently grounded on what Kant calls ―Erkenntnis‖. (shrink)

We set forth what one may call a “constitutional reading” in opposition to the traditional widespread “constructivist reading” of the object of cognition. In the light of the so-called one-object view reading of Transcendental idealism, the object of cognition is nothing but the object that exists in itself insofar as it appears to our cognitive apparatus. The object exists mind-independently, while our cognition of the same object must be mind-independent. The constructivist reading mistakes the epistemological problem of how we come (...) to cognize mind-dependently that what we represent (“the constitutional view”) are mind-independent objects with the Berkelian ontological problem of how we construct objects out of an undifferentiated, unstructured manifold (the constructivist view) My diagnosis is as follows. The first reason is the traditional “two-worlds view” reading of Kant’s idealism. If we take what exists in itself and the object of cognition as distinct things, then we must conclude that the object of cognition is a mind-dependent construction. Constructivist readers mistake the mind-dependent nature of our human cognition of objects for the putative mind-dependent nature of the known object. The second reason is overintellectualization. Constructivist readers mistake the objectifying syntheses of the imagination, below the threshold of self-consciousness, for cognitive conceptual operations by means of which we cognize (erkennen) the objects of cognition, or so I shall argue. (shrink)

This paper aims to present a novel reading of Kantian idealism. In want of a better name, I call my interpretation “one-object-plus-epistemic phenomenalism.” I partially endorse Allison’s celebrated position, namely his rejection of metaphysical world-dualism. Yet, I reject Allison’s deflationary two-aspect view. I argue that Kantian idealism is also metaphysically committed to an ontological noumenalism (one-object), namely the claim that the ultimate nature of reality is made up of unknown things in themselves (substantia noumena). Natural sciences can only reveal the (...) relational/structural properties of things as they appear rather than the intrinsic properties of substantia noumena in the negative sense. My anti-deflationary reading is similar to Allais’s, and my agnostic monism to Hanna’s. However, against both, I hold that appearances are not the accusative objects of our sensible representations but rather, as Kant repeatedly states: “mere representations.” The accusative objects of our senses are substantia noumena in the negative sense. Moreover, my view is also similar to Langton’s. Again, against Langton, I hold that appearances are not merely relational properties of substantia noumena but the way that such substantia noumena exist inside our mind as “mere representations.” In this regard, I also partially endorse Guyer’s and Van Cleve’s phenomenalist reading because these substantia noumena in the negative sense can only be cognized mind-dependently, namely as appearances. However, against Guyer and Van Cleve, I hold that the phenomenalist side of Kantian idealism is purely epistemological rather than ontological: what Kant calls the necessary unity of representations according to categories is not a logical construction of objects out of representations, that is, an ontological reduction of noumena to appearances, but rather the way we sense-independently cognize mind-independent noumena. (shrink)

I argue for a new reading of Pyrrhonian beliefs inspired by representationalism (the content view) in recent philosophy of mind. I shall argue that there are two senses of acceptance or “acquiescence in something” (eudokein tini pragmati) rather than two senses of belief (doxa). For this reason, we can maintain along with Sextus that the Pyrrhonian skeptic behaves intentionally and can live his life in society adoxastôs without any proper beliefs whatsoever. However, the skeptical sense of acceptance is not the (...) mere avowal of a feeling or a mere report or register of the way things appear to me. I want to suggest the following reading. Acceptance without belief is an anti-predicative passive and involuntary acceptance of content by means of an indicator function (pathos or phainomenon) in virtue of the past regular connection between the representational content and the indicator function. The pathos (as a present state of mind) or the phainomenon “means” (that is, indicates) passively and involuntarily a content in virtue of its regular connection with this content through time. For example, the Pyrrhonian’s acceptance of the content of the sentence “there is fire” is based on the fact that he here and now perceives smoke and that pathos passively and involuntarily means (represents) the content that there is fire. There is no judgment involved at all: even when the skeptic suspends his judgment (epochê) about whether there is fire, his pathos makes him involuntarily accept that there is fire. That is what Sextus, following the Stoic tradition, calls “commemorative signs” (hupomnêstikon) in opposition to the so-called “indicative signs” (endeiktikon), namely the “internal assertion” of the content as a belief (as a propositional attitude). As I do not believe that any reasonable reading of the skeptical acceptance may be defended only by means of philological analysis and by the research of the huge doxography, my arguments deliberately mix historical evidence with reasons of a systematic nature. In this way, the defense of my alternative reading is abductive, namely as the inference to the best explanation. (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)

The focus of this paper is Cassirer’s Neo-Kantian reading of Kant’s conception of unity of space. Cassirer’s neo-Kantian reading is largely in conformity with the mainstream of intellectualist Kant-scholars, which is unsurprising, given his own intellectualist view of space and perception and his rejection of the existence of a ‘merely sensory consciousness’ as a ‘formless mass of impression’. I argue against Cassirer’s reading by relying on a Kantian distinction first recognized by Heinrich Rickert, a neo-Kantian from the Southwest school, between (...) Kenntnis (roughly knowledge by acquaintance) and Erkenntnis (roughly propositional knowledge). Correspondingly, I claim that concepts and categories are conditions for Erkenntnis of objects as such, namely for thinking of and apprehending the pre-existing unity as an object, rather than for the ‘constitution’ of this very unity. (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)

Godfrey-Smith advocates for linking deception in sender-receiver games to the existence of undermining signals. I present games in which deceptive signals can be arbitrarily frequent, without this undermining information transfer between sender and receiver.

La Educación Sensible es ayuda para que el “yo” habite en su “hogar interior” y crezca hacia su “apoteosis original” en el “nosotros”, donde se hace cocreador de belleza con libertad, sabiduría y amor. La educación sensible es pedagogía no invasiva pero radicalmente exigente para que la persona acepte desplegar su versión original y vivir con gozo en un “nosotros-maduro”. No es una pedagogía que protege a los sensibles; es una educación que atiende a todas las personas, independientemente de su (...) grado de sensibilidad: alta, media o baja (como intervalos comunes que se manejan en las ciencias de la sensibilidad). La sensibilidad personal es capaz de captar la propia originalidad y disponerse para interpretarla en los dos sentidos: comprenderla y actuarla. La sensibilidad humana necesita la educación sensible para que sea punto de partida de un desarrollo personal auténtico y pleno de la propia originalidad que es alterada y altera la de los “yoes” con quien habita la existencia formando “nosotros”, que a su vez, se impactan con interdependencia. Además de al “yo”, la educación sensible se orienta al “nosotros”, que será tanto más “sensible” en la medida que altera la intimidad de la persona con “amor-maduro” que es fuente de gozo, a pesar de los pesares. Sin educación sensible la vida daña y cuanta mayor es la sensibilidad de la persona, mayor es el daño, pero también es mayor la capacidad para detectarlo cicatrizarlo y ayudar a otros con educación sensible. Así, la educación sensible se manifiesta de tres maneras: como acción sanadora de la persona, destrucción creadora para evolucionar en el tiempo, como la crisálida se transforma en mariposa, y acción desarrollante del propio potencial con originalidad. (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 paper addresses Fichte’s puzzle of self-consciousness. I propose a new reading of “Fichte’s original insight,” inspired by Pareyson’s general reading, which I call here the “Fichtean metaphysical turn in transcendental philosophy.” Against the mainstream view in Fichte’s scholarship, I argue that Fichte’s and Kant’s views do not concur regard- ing the primary reference of the “I”, namely spontaneous agency in thinking, which Fichte calls “Tathandlung”. Yet, their views do, in fact, concur when Fichte claims that this spontaneous agency in (...) thinking is the “essence” or the underlying nature of the self, which Kant denies. Regarding this, I take the side of Fichte. But how is Fichte’s original insight supposed to solve the puzzle of self-consciousness? At that transcendental level, the puzzle does not arise because there is no need for self-identification in the first place. Transcendental self-knowledge results from the sui generis intellectual Selbstanschauung that everyone has of oneself as sheer spontaneous agency in thinking. But at the empirical level, the puzzle does not arise either and for the same reason. Reference to the embodied self dispenses with any self-identification because it is based on the fundamental metaphysical relation everybody has to their own body, namely identity. (shrink)

La presente investigación tiene como propósito general asumir un posicionamiento filosófico en clave histórico-social y de tipo anti-relativista para analizar el desarrollo histórico de la matemática, el cual aplicaremos a un caso en particular: la matemática del antiguo Egipto. Para ello se discutirán y criticarán, en primera instancia, determinadas posiciones filosóficas afines al cuasi-empirismo en matemática que, siendo relativistas, permitirán delinear nuestro propio posicionamiento en contraste: la existencia de una «matemática situada». Esta categoría filosófica tendrá como sustento teórico la noción (...) de conocimiento situado. Además, las implicaciones de esta son, según sostenemos, tanto filosóficas como historiográficas, ya que servirá para analizar las características del corpus de problemas matemáticos egipcios registrados en los diversos papiros matemáticos. En particular, haremos referencia a las expresiones lingüísticas egipcias para denotar las diversas operaciones aritméticas y el carácter algorítmico de los problemas, así como también la cuestión epistemológica de la empiria y cómo la perspectiva situada permite superar la dicotomía entre una matemática pura y otra aplicada, que consideramos no es ubicua para abordar la interpretación de la práctica matemática del antiguo país del Nilo. Palabras clave: Filosofía de la matemática, Anti-relativismo, Matemática situada, Extrañeza del pasado, Matemática egipcia. (shrink)

Legal theorists seek to persuade other jurists of certain theories: Textualism or purposivism; formalism or realism; natural law theory or positivism; prison reform or abolition; universal or particular human rights? Despite voluminous literature about these debates, tremendous uncertainty remains about which views experts endorse. This Article presents the first-ever empirical study of American law professors about legal theory questions. A novel dataset of over six hundred law professors reveals expert consensus and dissensus about dozens of longstanding legal theory debates. -/- (...) Law professors also debate questions about the nature of the legal academy. Descriptively, which subjects (e.g. constitutional law) and methods (e.g. law & economics) are most central within the legal academy today? And prescriptively, should today’s legal academy prioritize additional areas (e.g. legislation) or methods (e.g. critical race theory)? There is great interest in these questions but no empirical dataset of experts’ views; this results in uncertainty about which views experts endorse. This Article’s empirical study also clarifies these questions, documenting law professors’ evaluation of over one-hundred areas of law. -/- The legal theory and legal academy findings support implications for legal scholarship, education, and practice. Clearly, debates about law and the legal academy’s evolution should not be settled by a survey. Nevertheless, insofar as law professors are experts about these issues, it is instructive to discover and carefully examine what views those experts hold, so as to help determine which views are most likely to be true and how the legal academy ought to develop. (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)

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.

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)

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.

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)