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)

Precision medicine and molecular systems medicine (MSM) are highly utilized and successful approaches to improve understanding, diagnosis, and treatment of many diseases from bench-to-bedside. Especially in the COVID-19 pandemic, molecular techniques and biotechnological innovation have proven to be of utmost importance for rapid developments in disease diagnostics and treatment, including DNA and RNA sequencing technology, treatment with drugs and natural products and vaccine development. The COVID-19 crisis, however, has also demonstrated the need for systemic thinking and transdisciplinarity and the limits (...) of MSM: the neglect of the bio-psycho-social systemic nature of humans and their context as the object of individual therapeutic and population-oriented interventions. COVID-19 illustrates how a medical problem requires a transdisciplinary approach in epidemiology, pathology, internal medicine, public health, environmental medicine, and socio-economic modeling. Regarding the need for conceptual integration of these different kinds of knowledge we suggest the application of general system theory (GST). This approach endorses an organism-centered view on health and disease, which according to Ludwig von Bertalanffy who was the founder of GST, we call Organismal Systems Medicine (OSM). We argue that systems science offers wider applications in the field of pathology and can contribute to an integrative systems medicine by (i) integration of evidence across functional and structural differentially scaled subsystems, (ii) conceptualization of complex multilevel systems, and (iii) suggesting mechanisms and non-linear relationships underlying the observed phenomena. We underline these points with a proposal on multi-level systems pathology including neurophysiology, endocrinology, immune system, genetics, and general metabolism. An integration of these areas is necessary to understand excess mortality rates and polypharmacological treatments. In the pandemic era this multi-level systems pathology is most important to assess potential vaccines, their effectiveness, short-, and long-time adverse effects. We further argue that these conceptual frameworks are not only valid in the COVID-19 era but also important to be integrated in a medicinal curriculum. (shrink)

How do opinion diversity and belief polarisation affect epistemic group decision-making, particularly if decisions must be made without delay and on the basis of permissive evidence? In an agent-based model, we track the consistency of group opinions aggregated through sentence-wise majority voting. Simulations on the model reveal that high opinion diversity, but not polarisation, incurs a significant inconsistency risk. These results indicate that epistemic group decisions based on permissive evidence can be particularly difficult for diverse groups. The results also improve (...) our understanding of what can reasonably be expected of expert groups, and where expert advice might have limits. (shrink)

As the world population is rapidly aging, stakeholders must address the care of the elderly with great concern. Also, loss of dignity is often associated with aging due to dementia, mobility problems and diminished functional autonomy. However, dignity is a polysemic term that is deemed useless by some ethicists. To counter this claim, we propose four concepts to define it better and make use accurately of this notion. These are human dignity, dignity of identity, dignities of excellence and attributed dignities. (...) Finally, we explain the importance of solicitude and human dignity in the care of the elderly. This will ensure the respect, friendship and dignity of the elderly in providing geriatric ethical care. (shrink)

The article analyses the problems and possibilities in saying that Judaism, Christianity and Islam ‹have the same God›.

Amongst the many social factors that impact upon children, race is arguably one of the largest. Race is an ever-present social category that governs many elements of a child’s interaction with others, and especially for racial minority children it exerts a deep influence on their understanding of themselves. In this chapter, we shall begin by examining what the concept of race really amounts to, emphasizing its status as a socially constructed concept, before examining in the following section how children first (...) come to recognize the existence of race, and to understand their own racial identity.We will then look at two important areas that illustrate the profound impact that the social presence of race and the child’s developing understanding of racial identity have upon the social conditions of many children. First, we will examine how race and childhood intersect in matters of educational opportunity and achievement, before moving on to examine the issue of transracial adoption in the final section. (shrink)

Recently, Jennifer Saul (“Racial Figleaves, the Shifting Boundaries of the Permissible, and the Rise of Donald Trump”, 2017; “Racist and Sexist Figleaves”, 2021) has explored the use of what she calls “figleaves” in the discourse on race and gender. Following Saul, a figleaf is an utterance that, for some portion of the audience, blocks the conclusion that some other utterance, R, or the person who uttered R is racist or sexist. Such racial and gender figleaves are pernicious, says Saul, because, (...) among other things, they can shift the boundaries of what is deemed acceptable to think or say. This paper expands on Saul’s picture in a threefold way: It is first argued that appeals to statistics can function as figleaves. It is then argued that—as far as figleaves go—there is reason to believe that statistics-as-figleaves are especially pernicious. Finally, the paper explores some strategies for counterspeech. (shrink)

Relying on aesthetic testimony seems problematic. For instance, it seems problematic for me to simply believe or assert that The Velvet Underground's debut album The Velvet Underground and Nico (1964) is amazing solely because you have told me so, even though I know you to be an honest and competent music critic. But why? After all, there do not seem to be similar reservations regarding testimony from many other domains. In this paper, I will argue that relying on aesthetic testimony (...) seems problematic because we are attached to an ideal of aesthetic authenticity and feel that living up to this ideal is anathema to simply relying on aesthetic testimony. (shrink)

Philosophers have offered many arguments to explain why historical injustices require reparations. This paper raises an unnoticed challenge for almost all of them. Most theories of reparations attempt to meet two intuitions: (1) Reparations are owed for a past wrong and (2) the content of reparations must reflect the historical injustice. I argue that necessarily no monistic theory can meet both intuitions. I do this by showing that any theory that can meet intuition (1) necessarily cannot also meet intuition (2). (...) This result suggests that a theory of reparations must either sacrifice one of the two intuitions or be pluralist. I argue that we ought to prefer the pluralist over the monistic theory. I sketch the pluralist theory and I defend it by considering an objection about the way it can resolve conflicts. (shrink)

In Necessary Beings, Bob Hale addresses two questions: What is the source of necessity? What is the source of our knowledge of it? He offers novel responses to them in terms of the metaphysical notion of nature or, more familiarly, essence. In this paper, I address Hale’s response to the first question. My assessment is negative. I argue that his essentialist explanation of the source of necessity suffers from three significant shortcomings. First, Hale’s leading example of an essentialist explanation merely (...) asserts that the nature of an entity explains some necessity, but leaves unexplained how it does so. Second, his essentialist explanation of particular necessities introduces new necessities that remain unexplained. Third, Hale’s version of essentialism presupposes a controversial metaphysical theory of properties, for which he offers no defense. (shrink)

Many philosophers, following Williamson (The Philosophical Review 105(4): 489–523, 1996), Williamson (Knowledge and its Limits, Oxford, Oxford Univer- sity Press, 2000), subscribe to the constitutive rule account of assertion (CRAA). They hold that the activity of asserting is constituted by a single constitutive rule of assertion. However, in recent work, Maitra (in: Brown & Cappelen (ed). Assertion: new philosophical essays, Oxford, Oxford University Press, 2011), Johnson (Acta Analytica 33(1): 51–67, 2018), and Kelp and Simion (Synthese 197(1): 125–137, 2020a), Kelp and (...) Simion (in: Goldberg (ed) The Oxford Handbook of Assertion, Oxford, Oxford University Press, 2020b) aim to show that, for all the most popular versions of the constitutive rule of assertion proposed in the literature, asserting is not an activity constituted by a single constitutive rule and that therefore CRAA is very likely false. To reach this conclusion, they all present a version of what can be dubbed the engagement condition objection. That is, they each propose a necessary condition on engaging in rule-constituted activities. Then they argue that, for all the most popular versions of the constitutive rule of assertion proposed in the literature, one can make assertions without satisfying this condition. In response, I present a counterexample that shows that the proposed engagement conditions lead to coun- terintuitive results, and I propose an alternative that better captures our intuitions. Then I argue that this alternative engagement condition is compatible with all the most popular versions of the constitutive rule of assertion. (shrink)

Gerade weil das Bestehen auf Rangordnungen in der heutigen Gesellschaft anstößig und fremd wirkt, ist es lohnenswert, sich ihnen mit Nietzsche neu zu stellen, der sie als sein Problem bezeichnete. Er richtet sie gezielt gegen die Gleichheit, von der er befürchtet, ihr Anspruch auf Universalität verunmögliche Individualität, Anders-Sein und damit auch alle Größe. Den moralischen Wert der Gleichheit kritisieren heißt nicht, sich von demokratischen Grundprinzipien oder Errungenschaften zu verabschieden. Geklärte Rangverhältnisse reduzieren Komplexität, vereinfachen die Kommunikation, machen Verhalten erwartbar und vereinfachen (...) so die Orientierung. So könnte gerade in den modernen Ausprägungen der Demokratie ein offenerer Umgang mit Rangordnungen zu ihrer Stärkung beitragen. Die Nietzsche-Forschung hat eine Beschäftigung mit dem Begriff bisher weitgehend vermieden. Alberts schließt diese Forschungslücke. Er geht persönlichen, philologischen und philosophischen Anhaltspunkten für Nietzsches Denken nach und untersucht die Perspektiven, die dabei auf verschiedenste Lebensbereiche wie Natur, Religion, Moral, Wissenschaft und Interindividualität eröffnet werden. (shrink)

Bilattices, introduced by Ginsberg as a uniform framework for inference in artificial intelligence, are algebraic structures that proved useful in many fields. In recent years, Arieli and Avron developed a logical system based on a class of bilattice-based matrices, called logical bilattices, and provided a Gentzen-style calculus for it. This logic is essentially an expansion of the well-known Belnap–Dunn four-valued logic to the standard language of bilattices. Our aim is to study Arieli and Avron’s logic from the perspective of abstract (...) algebraic logic . We introduce a Hilbert-style axiomatization in order to investigate the properties of the algebraic models of this logic, proving that every formula can be reduced to an equivalent normal form and that our axiomatization is complete w.r.t. Arieli and Avron’s semantics. In this way, we are able to classify this logic according to the criteria of AAL. We show, for instance, that it is non-protoalgebraic and non-self-extensional. We also characterize its Tarski congruence and the class of algebraic reducts of its reduced generalized models, which in the general theory of AAL is usually taken to be the algebraic counterpart of a sentential logic. This class turns out to be the variety generated by the smallest non-trivial bilattice, which is strictly contained in the class of algebraic reducts of logical bilattices. On the other hand, we prove that the class of algebraic reducts of reduced models of our logic is strictly included in the class of algebraic reducts of its reduced generalized models. Another interesting result obtained is that, as happens with some implicationless fragments of well-known logics, we can associate with our logic a Gentzen calculus which is algebraizable in the sense of Rebagliato and Verdú . We also prove some purely algebraic results concerning bilattices, for instance that the variety of distributive bilattices is generated by the smallest non-trivial bilattice. This result is based on an improvement of a theorem by Avron stating that every bounded interlaced bilattice is isomorphic to a certain product of two bounded lattices. We generalize it to the case of unbounded interlaced bilattices. (shrink)

This essay explores the nature of the most rudimentary form of empathy, interpersonal affective echoing, and attempts to give a cogent assessment of the roles it may play in human interactions. As an investigative background, it briefly sketches phenomenological findings with respect to feelings, to non-linguistic cognition, and to the analogical apperception of others. It then offers a phenomenological account of the basic structures of the experience of echoing another person’s feelings in a face-to-face situation. It also notes how echoing (...) differs from analogical apperception (while presupposing the latter), as well as from having one’s usual feelings and emotions. The relationship of echoing to sharing, caring, and morality is also briefly discussed. (shrink)

In medical ethics, business ethics, and some branches of political philosophy (multi-culturalism, issues of just allocation, and equitable distribution) the literature increasingly combines insights from ethics and the social sciences. Some authors in medical ethics even speak of a new phase in the history of ethics, hailing "empirical ethics" as a logical next step in the development of practical ethics after the turn to "applied ethics." The name empirical ethics is ill-chosen because of its associations with "descriptive ethics." Unlike descriptive (...) ethics, however, empirical ethics aims to be both descriptive and normative. The first question on which I focus is what kind of empirical research is used by empirical ethics and for which purposes. I argue that the ultimate aim of all empirical ethics is to improve the context-sensitivity of ethics. The second question is whether empirical ethics is essentially connected with specific positions in meta-ethics. I show that in some kinds of meta-ethical theories, which I categorize as broad contextualist theories, there is an intrinsic need for connecting normative ethics with empirical social research. But context-sensitivity is a goal that can be aimed for from any meta-ethical position. (shrink)

There is an ongoing debate in animalethics on the meaning and scope of animalwelfare. In certain broader views, leading anatural life through the development of naturalcapabilities is also headed under the conceptof animal welfare. I argue that a concern forthe development of natural capabilities of ananimal such as expressed when living freelyshould be distinguished from the preservationof the naturalness of its behavior andappearance. However, it is not always clearwhere a plea for natural living changes overinto a plea for the preservation (...) of theirnaturalness or wildness. In the first part ofthis article, I examine to what extent theconcerns for natural living meet ``theexperience requirement.'' I conclude that someof these concerns go beyond welfare. In thesecond part of the article. I ask whether wehave moral reasons to respect concernsfor the naturalness of an animal's living thattranscend its welfare. I argue that the moralrelevance of such considerations can be graspedwhen we see animals as entities bearingnon-moral intrinsic values. In my view the``natural'' appearance and behavior of an animalmay embody intrinsic values. Caring for ananimal's naturalness should then be understoodas caring for such intrinsic values. Intrinsicvalues provide moral reasons for action iffthey are seen as constitutive of the good lifefor humans. I conclude by reinterpreting,within the framework of a perfectionist ethicaltheory, the notion of indirect dutiesregarding animals, which go beyond andsupplement the direct duties towardsanimals. (shrink)

In the chapter alarmist ideas that exponential development of technologies is an exceptional source of the crisis of modern civilization are considered and reasonably questioned. The role of tools in the establishment of human and technologies is underlined, including information and computative processes in the formation of the modern culture. The role of information processes in nature and culture is explored, which enables us to make a conclusion that technologies including artificial intelligence and intelligent robotics are a compile and an (...) integral part of a human being. The role of algorithms as a deep basis of crisis events due to the new opportunities of a dialogue between the machine and the human and the Transition of information without a cultural context in human-machine system is justified. Various ways to overcome crisis events whilst human transformation are offered. Firstly it is the augmentation of human possibilities with the help of exoprosthetic devices or exocortexes which broaden physical and cognitive capabilities. Secondly it is the transformation of technologies basing on deep social and humanitarian research which demand expert answers to vital questions of using the developing technologies. The use of technologies is analyzed from the position of postneoclassic rationality and in this framework an attempt is taken to use postturing methodology which was used before on the example of intellectual robitics. There is also shown an opportunity to broaden the sphere of using this postturing methodology in the other kinds of technology, because gradually the ingress of artificial intelligence into all the tools of human activity will make any tool "an intellectual robot". The use of postturing methodology will enable to break a fundamental barrier which divides a human and a machine which will make the interaction of human and machine more effective and less antagonistic. To the further discussion a theological ground of technogenic civilization as creating an opportunity for life to overcome biological barriers and watch the change of cosmic scale is brought. Such ground enables to save the main achievement of the mankind - the culture. (shrink)

Some people wrongly believe that A. Turing’s works that underlie all modern computer science never discussed “physical” robots. This is not so, since Turing did speak about such machines, though making a reservation that this discussion was still premature. In particular, in his 1948 report [8], he suggested that a physical intelligent machine equipped with motors, cameras and loudspeakers, when wandering through the fields of England, would present “the danger to the ordinary citizen would be serious.” [8, ]. Due to (...) this imperfection of technology in the field of knowledge that we now call robotics, the methodology that he proposed was based on human speech, or rather on text. Other natural human skills were too difficult to implement, while the exchange of cues via written messages was much more accessible for engineering implementation in Turing’s time. Nevertheless, since then, the progress of computer technology has taken forms that the founder of artificial intelligence could not have foreseen. (shrink)

Pragmatist approaches to truth have often been judged in light of a caricature of William James’ claim that, “the ‘true’ is only the expedient in our way of thinking”. This unfortunate caricature, where truth is claimed to be ‘whatever it’s useful to believe’, means pragmatist theories of truth are generally seen as non-starters, or unworthy of serious attention. And even leaving aside stalking-horse versions of classical pragmatism, there is also a view that whatever contemporary pragmatists have been doing with ‘truth’ (...) it bears little resemblance to the projects that concern analytic philosophers. For example, Rorty’s frequent exhortations for pragmatists to turn their back on truth and look... (shrink)

Social entrepreneurship is usually understood as an economic activity which focuses at social values, goals, and investments that generates surpluses for social entrepreneurs as individuals, groups, and startups who are working for the benefit of communities, instead of strictly focusing mainly at the financial profit, economic values, and the benefit generated for shareholders or owners. Social entrepreneurship combines the production of goods, services, and knowledge in order to achieve both social and economic goals and allow for solidarity building. From a (...) broader perspective, entities that are focused on social entrepreneurship are identified as parts of the social and solidarity economy. These are, for example, social enterprises, cooperatives, mutual organizations, self-help groups, charities, unions, fair trade companies, community enterprises, and time banks. Social innovation is a key element of social entrepreneurship. Social innovation is usually understood as new strategies, concepts, products, services, and organizational forms that allow for the satisfaction of needs. Such innovations are created in particular in the contact areas of various sectors of the social system. For example, these are spaces between the public sector, the private sector, and civil society. These innovations not only allow the solving of problems but also extend possibilities for public action. (shrink)

As indicated in the title this article is a brief description of the body, soul and spirit of man, based on the divine revelations received by the Austrian mystic Jakob Lorber between 1840 and 1864. While it is common knowledge that man has a body and a soul, very little is known about the spirit in man which is the primary source of knowledge and power, penetrating both the soul and body.

Felix Maschewski und Anna-Verena Nosthoff untersuchen Facebook als Beispiel einer besonders eindringlichen Form „kybernetischer Gouvernementalität“ und nehmen das „soziale“ Netzwerk dazu als psychopolitisches System von Affekten und Aktivierungen in den Blick. Dabei analysieren sie v.a. die zugrundeliegende »Sozialphysik des Anstoßes« und ihre implizite kybernetische Logik, um den so forcierten (behavioristischen) Subjektbegriff zu beleuchten. In diesem Konnex wird nicht nur dargestellt, wie das gouvernementale Unternehmen qua feedbacklogischer Programmatik die Entscheidungen, das Verhalten oder gar die Wünsche seiner Nutzer_innen antizipiert und auf (...) diese Weise indirekt ‚regiert‘, sondern auch, welche Konsequenzen die konsumtive Plattformlogik für das Politische selbst zeitigt. Letztlich zeigt sich, dass digitale Apparaturen entgegen dem insb. im Silicon Valley populären Narrativ von einer technologischen Neutralität keineswegs bloße Mittel, sondern zuallererst Machtinstrumente sind. (shrink)

In this paper, I spell out a new challenge for epistemic internalism that emerges out of the current debate on testimonial justification. Based on recent developments within this debate, one might argue as follows: Epistemic internalism can’t account for the justification of our testimony-based beliefs, because (1) we should conceive of testimonial justification along anti-reductionist lines and (2) anti-reductionism is incompatible with epistemic internalism. In response to this challenge, I show that, despite initial appearances, epistemic internalism and anti-reductionism are compatible (...) after all. Therefore, being an anti-reductionist with regard to testimonial justification doesn’t force one to reject epistemic internalism. In order to argue for this result, I will make use of resources provided by speech act theory. (shrink)

In a previous work we studied, from the perspective ofAlgebraic Logic, the implicationless fragment of a logic introduced by O. Arieli and A. Avron using a class of bilattice-based logical matrices called logical bilattices. Here we complete this study by considering the Arieli-Avron logic in the full language, obtained by adding two implication connectives to the standard bilattice language. We prove that this logic is algebraizable and investigate its algebraic models, which turn out to be distributive bilattices with additional implication (...) operations. We axiomatize and state several results on these new classes of algebras, in particular representation theorems analogue to the well-known one for interlaced bilattices. (shrink)

Under which conditions are we epistemically justied to believe that what other people tell us is true? Traditionally, the answer has either been reductionist or anti-reductionist: Either our justication reduces to non-testimonial reasons, or we have a presumptive, though defeasible, right to believe what we are told. However, different cases pull in different directions. Intuitively, someone asking for the time is subject to different epistemic standards than a surgeon consulting a colleague before a dangerous operation. Following this line of thought, (...) this paper develops an account of testimonial justication that captures our reductionist as well as our anti-reductionist intuitions. It is argued that the speaker’s commitment to an epistemic norm, as well as the hearer’s understanding of that norm, gives the hearer a presumptive right to believe what she is told. However, this justication doesn’t apply to situations with high practical risks. Here, the hearer needs reductive reasons to believe that her interlocutor is especially qualied to give her the desired information. (shrink)

This paper is about what we call Deontically-flavored Nominal Constructions (DNCs) in English, such as "No ice cream" or "Dogs on leash only". DNCs are often perceived as commands and have been argued to be a type of non-canonical imperative, much like root infinitives in German or Russian. We argue instead that DNCs at their core are declaratives that cite a rule but can be used performatively in the right context. We propose that DNCs contain an elided deontic modal, i.e., (...) allowed, whose presence explains their distributional restrictions and interpretational properties. Among other things, we speculate on the licensing conditions of DNCs (the presence of 'only' or the negative determiner 'no'), suggesting that these are tied to the properties of discourses in which rules can be used naturally. (shrink)

Although the index is one of the best known features of Peirce's theory of signs there is little appreciation of Peirce's theory of the index amongst contemporary philosophers of language. Amongst Peirce scholars, the value placed on Peirce's account is greater, but is largely based on Thomas Goudge's paper, "Peirce's Index" (Goudge, 1965). Despite marking a crucial milestone in our comprehension of Peirce's theory, our understanding of indices and indexical reference has grown markedly over the last forty years. Time has (...) now come, then, to develop the work that Goudge began and to provide a full analysis of Peirce's account that makes its value to the philosophy of language clear. I undertake this enterprise here. (shrink)

This article offers comprehensive criticism of the Turing test and develops quality criteria for new artificial general intelligence (AGI) assessment tests. It is shown that the prerequisites A. Turing drew upon when reducing personality and human consciousness to “suitable branches of thought” re-flected the engineering level of his time. In fact, the Turing “imitation game” employed only symbolic communication and ignored the physical world. This paper suggests that by restricting thinking ability to symbolic systems alone Turing unknowingly constructed “the wall” (...) that excludes any possi-bility of transition from a complex observable phenomenon to an abstract image or concept. It is, therefore, sensible to factor in new requirements for AI (artificial intelligence) maturity assessment when approaching the Tu-ring test. Such AI must support all forms of communication with a human being, and it should be able to comprehend abstract images and specify con-cepts as well as participate in social practices. (shrink)

What is salvific community for Ignatius of Loyola? It is communion with Christ, a dynamic for which Ignatius used the expression ‹societies Jesu›. This wording has a revealing intertextuality. ‹Societas› is the Vulgate’s rendering of Pauline and Johannine koinōnia: «sharing in (Christ)». The NT overtones of the Ignatian experience of communion can be explored regarding a theology of relationship (person), of action (history) and of the Church (representation). Being a person is understood as being friend and servant, history as the (...) salvific kairos of Christ’s activity, and the Church as the communion representing him—each time a typically Christian conception deciding always for that option that cannot be generalised through abstraction. (shrink)

Albert Musschenga | : The central question of this article is, Are animals morally responsible for what they do? Answering this question requires a careful, step-by-step argument. In sections 1 and 2, I explain what morality is, and that having a morality means following moral rules or norms. In sections 3 and 4, I argue that some animals show not just regularities in their social behaviour, but can be rightly said to follow social norms. But are the norms they (...) follow also moral norms? In section 5, I contend, referring to the work of Shaun Nichols, that the basic moral competences or capacities are already present in nonhuman primates. Following moral rules or norms is more than just acting in accordance to these norms; it requires being motivated by moral rules. I explain, in section 6, referring to Mark Rowlands, that being capable of moral motivation does not require agency; being a moral subject is sufficient. Contrary to moral agents, moral subjects are not responsible for their behaviour. Stating that there are important similarities between animal moral behaviour and human, unconscious, automatic, habitual behaviour, I examine in section 7 whether humans are responsible for their habitual moral behaviour, and if they are, what then the grounds are for denying that moral animals are responsible for their behaviour. The answer is that humans are responsible for their habitual behaviour if they have the capacity for deliberate intervention. Although animals are capable of intervention in their habitual behaviour, they are not capable of deliberate intervention. | : La question centrale dans cet article est celle de savoir si les animaux sont moralement responsables de ce qu’ils font. Répondre à cette question nécessite une argumentation minutieuse et progressive. Dans les sections 1 et 2, j’explique ce qu’est la moralité, et qu’être doté de moralité signifie de se conformer à des règles ou à des normes morales. Dans les sections 3 et 4, je pose que certains animaux ne se contentent pas de montrer des régularités dans leur comportement social, mais qu’ils suivent aussi véritablement des normes sociales. Toutefois, les normes qu’ils suivent sont-elles morales? Dans la section 5, je prétends, en me référant aux travaux de Shaun Nichols, que les compétences ou capacités morales de base sont déjà présentes chez les primates non humains. Respecter des règles ou des normes morales est bien plus que d’agir conformément à ces normes; cela requiert d’être motivé par des règles morales. Dans la section 6, me référant à Mark Rowlands, j’explique que d’être capable de motivation morale ne nécessite pas d’être un agent moral; le fait d’être un sujet moral suffit. Contrairement aux agents moraux, les sujets moraux ne sont pas responsables de leur comportement. Affirmant qu’il y a de grandes similitudes entre le comportement moral d’un animal et le comportement inconscient, automatique et habituel d’un humain, je me penche, dans la section 7, sur la question de savoir si les humains sont responsables de leur comportement moral habituel, et si tel est le cas, sur quelle base on peut alors nier le fait que les animaux sont eux aussi responsables de leur comportement. La réponse à cette question est que les humains sont responsables de leur comportement habituel s’ils disposent d’une capacité d’intervention réfléchie, délibérée. Bien que les animaux soient capables d’intervenir dans leurs comportements habituels, ils ne peuvent le faire de manière réfléchie, délibérée. (shrink)

Computational models have shown how polarisation can rise among deliberating agents as they approximate epistemic rationality. This paper provides further support for the thesis that polarisation can rise under condition of epistemic rationality, but it does not depend on limitations that extant models rely on, such as memory restrictions or biased evaluation of other agents’ testimony. Instead, deliberation is modelled through agents’ purposeful introduction of arguments and their rational reactions to introductions of others. This process induces polarisation dynamics on its (...) own. A second result is that the effect size of polarisation dynamics correlates with particular types of argumentative behaviour. Polarisation effects can be soothed when agents take into account the opinions of others as premises, and they are amplified as agents fortify their own beliefs. These results underpin the relevance of argumentation as a factor in social-epistemic processes and indicate that rising issue polarisation is not a reliable indicator of epistemic shortcomings. (shrink)

In this paper, I critically assess different interpretations of Bohmian mechanics that are not committed to an ontology based on the wave function being an actual physical object that inhabits configuration space. More specifically, my aim is to explore the connection between the denial of configuration space realism and another interpretive debate that is specific to Bohmian mechanics: the quantum potential versus guidance approaches. Whereas defenders of the quantum potential approach to the theory claim that Bohmian mechanics is better formulated (...) as quasi-Newtonian, via the postulation of forces proportional to acceleration; advocates of the guidance approach defend the notion that the theory is essentially first-order and incorporates some concepts akin to those of Aristotelian physics. Here I analyze whether the desideratum of an interpretation of Bohmian mechanics that is both explanatorily adequate and not committed to configuration space realism favors one of these two approaches to the theory over the other. Contrary to some recent claims in the literature, I argue that the quasi-Newtonian approach based on the idea of a quantum potential does not come out the winner. (shrink)

Debates over the reality of race often rely on arguments about the connection between race and science—those who deny that race is real argue that there is no significant support from science for our ordinary race concepts; those who affirm that race is real argue that our ordinary race concepts are supported by scientific findings. However, there is arguably a more fundamental concern here: How should we define race concepts in the first place? The reason I claim that this definitional (...) question is more fundamental is that our handling of the underlying definitional problem often determines the scientific support our ordinary race concepts need, and importantly the likelihood of finding such support. In short, the defini- tional question, “How do we define race?” often undercuts the question of whether race is scientifically meaningful. (shrink)

With robots being introduced into caregiving, particularly for older persons, various ethical concerns are raised. Among them is the fear of replacing human caregiving. While ethical concepts like well-being, autonomy, and capabilities are often used to discuss these concerns, this paper brings forth the concept of social dignity to further develop guidelines concerning the use of robots in caregiving. By social dignity, we mean that a person’s perceived dignity changes in response to certain interactions and experiences with other persons. In (...) this paper, we will first present the concept of social dignity, and then identify a niche where robots can be used in caregiving in an ethical manner. Specifically, we will argue that, because some activities of daily living are performed in solitude to maintain dignity, a care recipient will usually prefer robotic assistance instead of human assistance for these activities. Secondly, we will describe how other philosophical concepts, which have been commonly used to judge robotic assistance in caregiving for the elderly so far, such as well-being, autonomy, and capabilities, are less useful in determining whether robotic assistance in caregiving is ethically problematic or not. To conclude, we will argue that social dignity offers an advantage to the other concepts, as it allows to ask the most pressing questions in caregiving. (shrink)

In this article I examine the consequences of the dominance of intuitive thinking in moral judging and deciding for the role of moral reasoning in moral education. I argue that evidence for the reliability of moral intuitions is lacking. We cannot determine when we can trust our intuitive moral judgements. Deliberate and critical reasoning is needed, but it cannot replace intuitive thinking. Following Robin Hogarth, I argue that intuitive judgements can be improved. The expertise model for moral development, proposed by (...) Hubert and Stuart Dreyfus, not only teaches us how we acquire intuitive moral judgements, it also shows the interconnectedness of intuitive thinking and deliberate reasoning. Analysing the expertise model in more detail, I show that it cannot do justice to the importance of reasoning skills. Reasoning skills are needed because we expect people to be able to argue for their standpoints. I conclude that moral education should not only aim at improving intuitive moral judgements, but also at acquiring reasoning skills. (shrink)

According to some proponents of an empirical medical ethics, medical ethics should take the experience, insights, and arguments of doctors and other medical practitioners as their point of departure. Medical practitioners are supposed to have ‘moral wisdom.’ In this view, the moral beliefs of medical practitioners have a special status. In sections I-IV, I discuss two possible defences of such a status. The first defence is based on the special status of the moral beliefs of the health professional as an (...) expert in medical ethics, and the second defence on the special status of the health professional’s moral beliefs as a practitioner. The first defence is built up around the opposition between experts and laypersons, while the second defence rests on the opposition between practitioners and theorists. In sections V and VI, I examine the interrelations between empirical ethics and anti-theory in ethics. Some regard empirical ethics as a laudable alternative to theory-driven normative ethics. I explore their objections against ethical theory and ethical theorising, and conclude that they are not valid. In the last section, VII, I discuss the role of ethicists in medical practice. (shrink)

In this paper I examine parallels between C.S. Peirce's most mature account of signs and contemporary philosophy of language. I do this by first introducing a summary of Peirce's final account of Signs. I then use that account of signs to reconstruct Peircian answers to two puzzles of reference: The Problem of Cognitive Significance, or Frege's Puzzle; and The Same-Saying Phenomenon for Indexicals. Finally, a comparison of these Peircian answers with both Fregean and Direct Referentialist approaches to the puzzles highlights (...) interesting parallels and important differences between Peirce's final account of signs, and the concepts used in analytic philosophy of language. (shrink)

People partaking in a conversation can add to the common ground of said conversation by performing different speech acts. That is, they can influence which propositions are presumed to be shared among them. In this paper, I am going to apply the common ground framework to the phenomenon of epistemic injustice. In doing so, I am going to focus on two kindsof speech acts: making assertions and asking certain kinds of questions. And I am going to look at three varieties (...) of epistemic injustice: testimonial injustice, inquiring injus-tice and interpretative injustice.I am going to argue that what all these varieties of epistemic injustice have in common is that they unfairly inhibit the speaker’s ability to add to the common ground in the way intended by her. This in turn negatively affects which con-versational roles a speaker can play in a given conversation. Based on these results, I am going to end by looking at some of the harms that epistemic injustice inflicts upon its victims. (shrink)

This article examines the various Liar paradoxes and their near kin, Grelling’s paradox and Gödel’s Incompleteness Theorem with its self-referential Gödel sentence. It finds the family of paradoxes to be generated by circular definition–whether of statements, predicates, or sentences–a manoeuvre that generates pseudo-statements afflicted with the Liar syndrome: semantic vacuity, semantic incoherence, and predicative catalepsy. Such statements, e.g., the self-referential Liar statement, are meaningless, and hence fail to say anything, a point that invalidates the reasoning on which the various paradoxes (...) rest. The seeming plausibility of the paradoxes is due to the fact that often the sentence used to make the pseudo-statement is ambiguous in that it may also be used to make a genuine statement about the pseudo-statement. Hence, if a formal system is to avoid ambiguity and consequent paradox and contradiction, it must distinguish between the two statements the sentence may be used to make. Gödel’s Theorem presents a further complication in that the self-reference involved is sentential rather than statemental. Nevertheless, on the intended interpretation of the system as a formalization of arithmetic, the self-referential Gödel sentence can only be an ambiguous statement, one that is both a pseudo-statement and its genuine double. Consequently, the conclusions commonly drawn from Gödel’s theorem must be deemed unwarranted. Arithmetic might well be formalized in a proper system that either excludes circular definition or introduces disambiguators. (shrink)

This paper addresses a family of issues surrounding the biological phenomenon of resistance and its representation in realist ontologies. The treatments of resistance terms in various existing ontologies are examined and found to be either overly narrow, internally inconsistent, or otherwise problematic. We propose a more coherent characterization of resistance in terms of what we shall call blocking dispositions, which are collections of mutually coordinated dispositions which are of such a sort that they cannot undergo simultaneous realization within a single (...) bearer. A definition of ‘protective resistance’ is proposed for use in the Infectious Disease Ontology (IDO) and we show how this definition can be used to characterize the antibiotic resistance in Methicillin-Resistant Staphylococcus aureus (MRSA). The ontological relations between entities in our MRSA case study are used alongside a series of logical inference rules to illustrate logical reasoning about resistance. A description logic representation of blocking dispositions is also provided. We demonstrate that our characterization of resistance is sufficiently general to cover two other cases of resistance in the infectious disease domain involving HIV and malaria. (shrink)

Este artigo discute a relação entre a questão das inferências nos contextos de justificação em ciência, nomeadamente o que ficou conhecido por “problema da indução” após Hume, em vista de esclarecer diferentes versões dos problemas de subdeterminação na filosofia da ciência.

Rationalists often argue that empiricism is incoherent and conclude, on that basis, that some knowledge is a priori. I contend that such arguments against empiricism cannot be parlayed into an argument in support of the a priori since rationalism is open to the same arguments. I go on to offer an alternative strategy. The leading idea is that, instead of offering a priori arguments against empiricism, rationalists should marshal empirical support for their position.

Der vorliegende Forschungsband möchte der kulturelle Bedeutung der Ersten Republik und deren Leistungen einen gebührenden Platz einräumen. Während sich nämlich Wiener Moderne und Weimarer Republik heute einer fortdauernden Aufmerksamkeit erfreuen, sind die Kultur der Ersten Republik und hier vorab des soziokulturellen Laboratoriums Wien – vermutlich schon in der Wahrnehmung vieler Zeitgenossinnen und Zeitgenossen unterbelichtet geblieben. Versammelt sind Beiträge zur Volksbildung im Roten Wien, zu Architektur und Stadtplanung, zu neuen Diskursen in der Psychoanalyse oder zur Neuorientierung Musils nach 1918, die später (...) zu seinem epochalen Roman »Der Mann ohne Eigenschaften« führen wird. Was ihn von anderen Publikationen unterscheidet, ist ein komparatistischer Aspekt, der etwa im Vergleich zwischen der Revolution 1918 in Wien und Österreich und der (kurzlebigen) Räterepublik in Ungarn aber auch in der Analyse der ungarischen Medien in der Zwischenkriegszeit zum Ausdruck kommt. Durch diese erhellenden Vergleiche fällt neues Licht auf die spezifische Besonderheit der Ersten Republik und der Situation im Roten Wien. (shrink)

This paper addresses the use of dispositions in the Infectious Disease Ontology (IDO). IDO is an ontology constructed according to the principles of the Open Biomedical Ontology (OBO) Foundry and uses the Basic Formal Ontology (BFO) as an upper ontology. After providing a brief introduction to disposition types in BFO and IDO, we discuss three general techniques for representing combinations of dispositions under the headings blocking dispositions, complementary dispositions, and collective dispositions. Motivating examples for each combination of dispositions is given (...) along with a specific use case in IDO. Description logic restrictions are used to formalize statements relating to these combinations. (shrink)

Definition of the problem The umbrella term “explicability” refers to the reduction of opacity of artificial intelligence (AI) systems. These efforts are challenging for medical AI applications because higher accuracy often comes at the cost of increased opacity. This entails ethical tensions because physicians and patients desire to trace how results are produced without compromising the performance of AI systems. The centrality of explicability within the informed consent process for medical AI systems compels an ethical reflection on the trade-offs. Which (...) levels of explicability are needed to obtain informed consent when utilizing medical AI? Arguments We proceed in five steps: First, we map the terms commonly associated with explicability as described in the ethics and computer science literature, i.e., disclosure, intelligibility, interpretability, and explainability. Second, we conduct a conceptual analysis of the ethical requirements for explicability when it comes to informed consent. Third, we distinguish hurdles for explicability in terms of epistemic and explanatory opacity. Fourth, this then allows to conclude the level of explicability physicians must reach and what patients can expect. In a final step, we show how the identified levels of explicability can technically be met from the perspective of computer science. Throughout our work, we take diagnostic AI systems in radiology as an example. Conclusion We determined four levels of explicability that need to be distinguished for ethically defensible informed consent processes and showed how developers of medical AI can technically meet these requirements. (shrink)

Christopher Hill contends that the metaphysical modalities can be reductively explained in terms of the subjunctive conditional and that this reductive explanation yields two tests for determining the metaphysical modality of a proposition. He goes on to argue that his reductive account of the metaphysical modalities in conjunction with his account of modal knowledge underwrites the further conclusion that conceivability does not provide a reliable test for metaphysical possibility. I argue (1) that Hill’s reductive explanation of the metaphysical modalities in (...) terms of the subjunctive conditional does not yield a reductive explanation of knowledge of metaphysical modality in terms of knowledge of subjunctive conditionals, and (2) that his account of modal knowledge is at odds with his contention that conceivability does not provide epistemic access to metaphysical possibility. (shrink)