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)

This essay contains an attempt to trace the evolution of the concept of wisdom as found in the thought of Aristotle and Aquinas in terms of how the philosophical concept of wisdom as an intellectual virtue is understood and used to express the theological concept of wisdom as a gift of the Holy Spirit. The main aim is to understand how Aquinas derived the concept of wisdom from Aristotle's metaphysics and developed it in his mysticism. This research is based on (...) a close study of Book Six of Aristotle's Nichomachean Ethics, the corresponding sections of Aquinas' Sententia libri Ethicorum and question forty-five of the second part of the second part of Aquinas' Summa Theologiae. The insights gained from the study are then used to decipher the theoretical meaning of Augustine's famous saying: "love and do what thou wilt" and to expound on the practical value of wisdom for religious leaders. (shrink)

In contemporary debates in religious epistemology, theistic philosophers provide differing responses to the evidentialist argument against religious beliefs. Plantinga’s strategy is to argue that evidence is not needed to justify religious beliefs while Swinburne’s strategy is to argue that religious beliefs can be justified by evidence. However, in Aquinas’ account of religious epistemology, he seems to employ both strategies. In his account of religious knowledge by faith, he argues that evidence is unnecessary for religious beliefs. But in his account of (...) religious knowledge by science, he argues that there is evidence for religious beliefs. In this paper, I argue that there is no real dichotomy between Plantinga’s and Swinburne’s responses to the evidentialist argument. From a Thomistic perspective, Reformed Epistemology and Natural Theology are different but compatible responses to Evidentialism. (shrink)

In this paper, I explore what gives collective testimony its epistemic credentials, through a critical discussion of three competing accounts of the epistemology of collective testimony. According to the first view, collective testimony inherits its epistemic credentials from the beliefs the testimony expresses— where this can be seen either as the beliefs of all or some of the group’s members, or as the beliefs of group itself. The second view denies any necessary connection to belief, claiming instead that the epistemic (...) credentials of collective testimony derive from the reliability or truth-conduciveness of the statement that expresses the testimony. Finally, the third view claims that the epistemic credentials of collective testimony derive from the fact that it involves undertaking a collective commitment to trustworthiness, which makes the group susceptible to rebuke and blame if its testimony is not trustworthy. I argue that this last account holds the most promise for preserving what is distinctive about testimonial knowledge while still underwriting a robust epistemology of collective testimony. (shrink)

Over the past few decades, Indigenous communities have successfully campaigned for greater inclusion in decision-making processes that directly affect their lands and livelihoods. As a result, two important participatory rights for Indigenous peoples have now been widely recognized: the right to consultation and the right to free, prior and informed consent (FPIC). Although these participatory rights are meant to empower the speech of these communities—to give them a proper say in the decisions that most affect them—we argue that the way (...) these rights have been implemented and interpreted sometimes has the opposite effect, of denying them a say or ‘silencing’ them. In support of this conclusion we draw on feminist speech act theory to identify practices of locutionary, illocutionary, and perlocutionary group silencing that arise in the context of consultation with Indigenous communities. (shrink)

Recent feminist philosophy of language has highlighted the ways that the speech of women can be unjustly impeded, because of the way their gender affects the uptake their speech receives. In this chapter, I explore how similar processes can undermine the speech of a different sort of speaker: Indigenous communities. This involves focusing on Indigeneity rather than gender as the salient social identity, and looking at the ways that group speech, rather than only individual speech, can be unjustly impeded. To (...) do this, I make use of the notion of ‘discursive injustice’ that has been developed by Quill Kukla, and, with reference to three case studies, show how discursive injustice can effectively derail the speech of Indigenous communities. (shrink)

Jennifer Lackey (2018) has developed an account of the primary form of group assertion, according to which groups assert when a suitably authorized spokesperson speaks for the group. In this paper I pose a challenge for Lackey's account, arguing that her account obscures the phenomenon of group silencing. This is because, in contrast to alternative approaches that view assertions (and speech acts generally) as social acts, Lackey's account implies that speakers can successfully assert regardless of how their utterances are taken (...) up by their audiences. What reflection on group silencing shows us, I argue, is that an adequate account of group assertion needs to find a place for audience uptake. (shrink)

According Philip Pettit, suitably organised groups not only possess ‘minds of their own’ but can also ‘make up their minds’ and 'speak for themselves'--where these two capacities enable them to perform as conversable subjects or 'persons'. In this paper I critically examine Pettit's case for group personhood. My first step is to reconstruct his account, explaining first how he understands the two capacities he considers central to personhood – the capacity to ‘make up one’s mind’, and the capacity to ‘speak (...) for oneself’ – before showing how he thinks these can be manifested in groups. With Pettit’s account duly reconstructed, I then turn to criticism, arguing that Pettit’s construal of making up one’s mind does not do proper justice to our first-personal self-understanding, nor to our characteristic interpersonal forms of engagement. This leads me, finally, to consider an alternative construal of ‘making up one’s mind’ and ‘speaking for oneself’ that is associated with the work of Richard Moran and whichargue, could usefully be exteextended to groups. (shrink)

In this paper, I set out a new Kantian approach to resolving conflicts between moral obligations for highly autonomous machine agents. First, I argue that efforts to build explicitly moral autonomous machine agents should focus on what Kant refers to as duties of right, which are duties that everyone could accept, rather than on duties of virtue (or “ethics”), which are subject to dispute in particular cases. “Moral” machines must first be rightful machines, I argue. I then show how this (...) shift in focus from ethics to a standard of public right resolves the conflicts in what is known as the "trolley problem" for autonomous ma-chine agents. Finally, I consider how a deontic logic suitable for capturing duties of right might meet Kant’s requirement that rightfully enforceable obligations be consistent in a system of equal freedom under universal law. (shrink)

In this paper, I set out some basic elements of a deontic logic with an implementation appropriate for handling conflicting legal obligations for purposes of programming autonomous machine agents. Kantian justice demands that the prescriptive system of enforceable public laws be consistent, yet statutes or case holdings may often describe legal obligations that contradict; moreover, even fundamental constitutional rights may come into conflict. I argue that a deontic logic of the law should not try to work around such conflicts but, (...) instead, identify and expose them so that the rights and duties that generate inconsistencies in public law can be explicitly qualified and the conflicts resolved. I then argue that a credulous, non-monotonic deontic logic can describe inconsistent legal obligations while meeting Kant’s demand for consistency in the prescriptive system of public law. Finally, I propose an implementation of this logic via a modified form of “answer set programming,” which I demonstrate with some simple examples. (shrink)

The central challenge of “machine ethics” is to build autonomous machine agents that act morally rightly. But how can we build autonomous machine agents that act morally rightly, given reasonable disputes over what is right and wrong in particular cases? In this chapter, I argue that Immanuel Kant’s political philosophy can provide an important part of the answer.

Moral abolitionists claim that morality ought to be abolished. According to one of their most prominent arguments, this is because making moral judgments renders people significantly less tolerant toward anyone who holds divergent views. In this paper we investigate the hypothesis that morality’s tolerance-decreasing effect only occurs if people are realists about moral issues, i.e., they interpret these issues as objectively grounded. We found support for this hypothesis (Studies 1 and 2). Yet, it also turned out that the intolerance associated (...) with realism is mediated by moral conviction and perceived consensus. People tend to feel more strongly about those moral issues they ground objectively and, in doing so, are more prone to display the vice of moral smugness toward those who disagree with them. The remedy for this that has been recommended is humility which we found (Study 3) is indeed related to reduced intolerance, in part by predicting a reduction in realism, but also in part through a direct connection to intolerance. These results put pressure on abolitionists’ “argument from intolerance.”. (shrink)

Introductory students regularly endorse naïve skepticism—unsupported or uncritical doubt about the existence and universality of truth—for a variety of reasons. Though some of the reasons for students’ skepticism can be traced back to the student—for example, a desire to avoid engaging with controversial material or a desire to avoid offense—naïve skepticism is also the result of how introductory courses are taught, deemphasizing truth to promote students’ abilities to develop basic disciplinary skills. While this strategy has a number of pedagogical benefits, (...) it prevents students in early stages of intellectual development from understanding truth as a threshold concept. Using philosophy as a case study, I argue that we can make progress against naïve skepticism by clearly discussing how metadisciplinary aims differ at the disciplinary and course levels in a way that is meaningful, reinforced, and accessible. (shrink)

The ontic conception of scientific explanation has been constructed and motivated on the basis of a putative lexical ambiguity in the term explanation. I raise a puzzle for this ambiguity claim, and then give a deflationary solution under which all ontically-rendered talk of explanation is merely elliptical; what it is elliptical for is a view of scientific explanation that altogether avoids the ontic conception. This result has revisionary consequences for New Mechanists and other philosophers of science, many of whom have (...) assimilated their conception of explanation to the ontic conception. (shrink)

What features will something have if it counts as an explanation? And will something count as an explanation if it has those features? In the second half of the 20th century, philosophers of science set for themselves the task of answering such questions, just as a priori conceptual analysis was generally falling out of favor. And as it did, most philosophers of science just moved on to more manageable questions about the varieties of explanation and discipline-specific scientific explanation. Often, such (...) shifts are sound strategies for problem-solving. But leaving fallow certain basic conceptual issues can also result in foundational debates. (shrink)

Politics, Philosophy & Economics, Volume 21, Issue 4, Page 347-371, November 2022. We sometimes have to choose between options that are seemingly incomparable insofar as they seem to be neither better than, worse than, nor equal to each other. This often happens when the available options are quite different from one another. For instance, consider a choice between prioritizing either criminal justice reform or healthcare reform as a public policy goal. Even after the relevant details of the goals and possible (...) reforms are filled in, it is plausible that neither goal is better than, worse than, nor equal to the other. Such seemingly incomparable options present a problem for rational choice since it is unclear how an agent might rationally choose between them. What we need are some principles to help govern rational choice when faced with seemingly incomparable options. I here present three such principles. While each principle is individually compelling, I show that they are jointly incompatible. I then argue that the correct response to this inconsistent triad is to reject the principle that rationally censures performing a sequence of choices one knows will result in a suboptimal outcome. The upshot is that when seeming incomparability is involved, an agent can money pump themselves without being less rational for it. (shrink)

I explore the possibility and rationality of interpersonal mechanisms of doxastic self-control, that is, ways in which individuals can make use of other people in order to get themselves to stick to their beliefs. I look, in particular, at two ways in which people can make interpersonal epistemic commitments, and thereby willingly undertake accountability to others, in order to get themselves to maintain their beliefs in the face of anticipated “epistemic temptations”. The first way is through the avowal of belief, (...) and the second is through the establishment of collective belief. I argue that both of these forms of interpersonal epistemic commitment can function as effective tools for doxastic self-control, and, moreover, that the control they facilitate should not be dismissed as irrational from an epistemic perspective. (shrink)

In this paper I critically discuss Miranda Fricker’s ‘trust-based’ view of collective testimony—that is, testimony that comes from a group speaker. At the heart of Fricker’s account is the idea that testimony involves an ‘interpersonal deal of trust’, to which the speaker contributes a commitment to ‘second-personal epistemic trustworthiness’. Appropriating Margaret Gilbert’s concept of joint commitment, Fricker suggests that groups too can make such commitments, and hence that they, like individuals, can ‘enter into the second-personal relations of trust that characterise (...) testimony’ (Fricker 2012: 272). I argue that this choice to appropriate Gilbert’s concept of joint commitment betrays a deep problem in Fricker’s account—a misconstrual of both the object and the subject(s) of the commitment a speaker makes in testifying. After developing this criticism, I outline an alternative way of construing the speaker’s commitment, which can be applied to both collective and individual testimony. (shrink)

For much of the first fifty years of its existence, analytic philosophy shunned discussions of normativity and ethics. Ethical statements were considered as pseudo-propositions, or as expressions of pro- or con-attitudes of minor theoretical significance. Nowadays, in contrast, prominent analytic philosophers pay close attention to normative problems. Here we focus our attention on the work of Searle, at the same time drawing out an important connection between Searle’s work and that of two other seminal figures in this development: H.L.A. Hart (...) and John Rawls. We show that all three thinkers tend to assume that there is but one type of normativity within the realm of social institutions – roughly, the sort of normativity that is involved in following the results of chess – and that they thereby neglect features that are of crucial significance for an adequate understanding of social reality. (shrink)

In this paper, I develop two philosophically suggestive arguments that the late Justice Stevens made in Citizens United against the idea that business corporations have free speech rights. First, (1) while business corporations conceived as real entities are capable of a thin agency conceptually sufficient for moral rights, I argue that they fail to clear important justificatory hurdles imposed by interest or choice theories of rights. Business corporations conceived as real entities lack an interest in their personal security; moreover, they (...) are incapable of exercising innate powers of choice. Second, (2) I argue that the structure and functionally individualized purpose of a business corporation—to increase value for its shareholders—undermines the implicit joint commitment necessary to derive corporate rights of free speech from non-operative shareholder-member rights. Since one cannot transfer innate moral rights such as free speech, any exercise of this right on behalf of another must be limited in scope. (shrink)

Psychoneural reductionists sometimes claim that sufficient amounts of lower-level explanatory achievement preclude further contributions from higher-level psychological research. Ostensibly, with nothing left to do, the effect of such preclusion on psychological explanation is extinction. Reductionist arguments for preclusion have recently involved a reorientation within the philosophical foundations of neuroscience---namely, away from the philosophical foundations and toward the neuroscience. In this chapter, I review a successful reductive explanation of an aspect of reward function in terms of dopaminergic operations of the mesocorticolimbic (...) system in order to demonstrate why preclusion/extinction claims are dubious. (shrink)

As much as assumptions about mechanisms and mechanistic explanation have deeply affected psychology, they have received disproportionately little analysis in philosophy. After a historical survey of the influences of mechanistic approaches to explanation of psychological phenomena, we specify the nature of mechanisms and mechanistic explanation. Contrary to some treatments of mechanistic explanation, we maintain that explanation is an epistemic activity that involves representing and reasoning about mechanisms. We discuss the manner in which mechanistic approaches serve to bridge levels rather than (...) reduce them, as well as the different ways in which mechanisms are discovered. Finally, we offer a more detailed example of an important psychological phenomenon for which mechanistic explanation has provided the main source of scientific understanding. (shrink)

A philosopher once wrote the following words:If I examine the PTOLOMAIC and COPERNICAN systems, I endeavour only, by my enquiries, to know the real situation of the planets; that is, in other words, I endeavour to give them, in my conception, the same relations, that they bear towards each other in the heavens. To this operation of the mind, therefore, there seems to be always a real, though often an unknown standard, in the nature of things; nor is truth or (...) falsehood variable by the various apprehensions of mankind. Though all human race should for ever conclude, that the sun moves, and the earth remains at rest, the sun stirs not an inch from his place for all these reasonings; and such conclusions are eternally false and erroneous. (shrink)

This manuscript discusses the author’s experience implementing a secularized version of Lectio Divina, a medieval monastic contemplative reading practice, in an introductory philosophy classroom. Following brief discussion of Lectio Divina’s history and a description of how the practice was modified for the classroom, I discuss three benefits (increased attention to cognitive and noncognitive reactions to the text, willingness to engage with the material in novel ways, and the opportunity to engage in independent disciplinary practice) and three potential challenges (the time (...) required, student engagement, and the practice’s perceived religiosity) arising from the exercise. Following this, I discuss potential modifications to the exercise that instructors may wish to consider, namely strategies for addressing students’ status as novice meditators, focusing textual selections on course materials, and having students engage in some aspects of the practice as homework. (shrink)

Traditional inflationary approaches that specify the nature of truth are attractive in certain ways; yet, while many of these theories successfully explain why propositions in certain domains of discourse are true, they fail to adequately specify the nature of truth because they run up against counterexamples when attempting to generalize across all domains. One popular consequence is skepticism about the efficaciousness of inflationary approaches altogether. Yet, by recognizing that the failure to explain the truth of disparate propositions often stems from (...) inflationary approaches' allegiance to alethic monism, pluralist approaches are able to avoid this explanatory inadequacy and the resulting skepticism, though at the cost of inviting other conceptual difficulties. A novel approach, alethic functionalism, attempts to circumvent the problems faced by pluralist approaches while preserving their main insights. Unfortunately, it too generates additional problems---namely, with its suspect appropriation of the multiple realizability paradigm and its platitude-based strategy---that need to be dissolved before it can constitute an adequate inflationary approach to the nature of truth. (shrink)

This essay presents deductive arguments to an introductory-level audience via a discussion of Aristotle's three types of rhetoric, the goals of and differences between deductive and non-deductive arguments, and the major features of deductive arguments (e.g., validity and soundness).

Each annual Ontology Summit initiative makes a statement appropriate to each Summit’s theme as part of our general advocacy designed to bring ontology science and engineering into the mainstream. The theme this year is "Towards an Open Ontology Repository". This communiqué represents the joint position of those who were engaged in the year's summit discourse on an Open Ontology Repository (OOR) and of those who endorse below. In this discussion, we have agreed that an "ontology repository is a facility where (...) ontologies and related information artifacts can be stored, retrieved and managed." -/- We believe in the promise of semantic technologies based on logic, databases and the Semantic Web, a Web of exposed data and of interpretations of that data (i.e., of semantics), using common standards. Such technologies enable distinguishable, computable, reusable, and sharable meaning of Web and other artifacts, including data, documents, and services. We also believe that making that vision a reality requires additional supporting resources and these resources should be open, extensible, and provide common services over the ontologies. (shrink)

One type of argument to sceptical paradox proceeds by making a case that a certain kind of metaphysically “heavyweight or “cornerstone” proposition is beyond all possible evidence and hence may not be known or justifiably believed. Crispin Wright has argued that we can concede that our acceptance of these propositions is evidentially risky and still remain rationally entitled to those of our ordinary knowledge claims that are seemingly threatened by that concession. A problem for Wright’s proposal is the (...) so-called Leaching worry: if we are merely rationally entitled to accept the cornerstones without evidence, how can we achieve evidence-based knowledge of the multitude of quotidian propositions that we think we know, which require the cornerstones to be true? This paper presents a rigorous, novel explication of this worry within a Bayesian framework, and offers the Entitlement theorist two distinct responses. (shrink)

When talking about truth, we ordinarily take ourselves to be talking about one-and-the-same thing. Alethic monists suggest that theorizing about truth ought to begin with this default or pre-reflective stance, and, subsequently, parlay it into a set of theoretical principles that are aptly summarized by the thesis that truth is one. Foremost among them is the invariance principle.

Functionalists about truth employ Ramsification to produce an implicit definition of the theoretical term _true_, but doing so requires determining that the theory introducing that term is itself true. A variety of putative dissolutions to this problem of epistemic circularity are shown to be unsatisfactory. One solution is offered on functionalists' behalf, though it has the upshot that they must tread on their anti-pluralist commitments.

Minimalists about truth contend that traditional inflationary theories systematically fail to explain certain facts about truth, and that this failure licenses a ‘reversal of explanatory direction’. Once reversed, they purport that their own minimal theory adequately explains all of the facts involving truth. But minimalists’ main objection to inflationism seems to misfire, and the subsequent reversal of explanatory direction, if it can be made sense of, leaves minimalism in no better explanatory position; and even if the objection were serviceable and (...) the reversal legitimate, minimalists’ adequacy thesis is still implausible. (shrink)

A restricted-use mobile device policy for introductory philosophy classrooms is presented and defended. The policy allows students to use devices only during open periods announced by the professor and is based on recent empirical findings on the effects of in-class mobile device use. These results suggest devices are generally detrimental to student learning, though they have targeted benefits for specific tasks. The policy is defended via a discussion of the ethical considerations surrounding device use, a discussion of the policy’s benefits, (...) and responses to potential objections. Avenues for future research are suggested at the conclusion of the discussion. (shrink)

Progressive stacking is a strategy for prioritizing in-class contributions that allows marginalized students to speak before non-marginalized students. I argue that this strategy is both pedagogically and ethically defensible. Pedagogically, it provides benefits to all students (e.g., expanded in-class discourse) while providing special benefits (e.g., increased self-efficacy) to marginalized students, helping to address historic educational inequalities. Ethically, I argue that neither marginalized nor non-marginalized students are wronged by such a policy. First, I present a strategy for self-disclosure that reduces the (...) risk of inadvertent, unwanted disclosure while respecting marginalized student autonomy in a manner analogous to accommodations provided under the Americans with Disabilities Act. Second, I argue that non-marginalized students are not wronged because such students are not silenced during discussion and because non-marginalized students benefit from the prioritization of marginalized voices. (shrink)

Pluralists maintain that there is more than one truth property in virtue of which bearers are true. Unfortunately, it is not yet clear how they diagnose the liar paradox or what resources they have available to treat it. This chapter considers one recent attempt by Cotnoir (2013b) to treat the Liar. It argues that pluralists should reject the version of pluralism that Cotnoir assumes, discourse pluralism, in favor of a more naturalized approach to truth predication in real languages, which should (...) be a desideratum on any successful pluralist conception. Appealing to determination pluralism instead, which focuses on truth properties, it then proposes an alternative treatment to the Liar that shows liar sentences to be undecidable. (shrink)

I argue that campus closures and shifts to online instruction in the early stages of the COVID-19 pandemic created an obligation to offer courses asynchronously. This is because some students could not have reasonably foreseen circumstances making continued synchronous participation impossible. Offering synchronous participation options to students who could continue to participate thusly would have been unfair to students who could not participate synchronously. I also discuss why ex post facto consideration of this decision is warranted, noting that similar actions (...) may be necessary in the future and that other tough pedagogical cases share important similarities with this case. (shrink)

Embodied Cognition is the kind of view that is all trees, no forest. Mounting experimental evidence gives it momentum in fleshing out the theoretical problems inherent in Cognitivists’ separation of mind and body. But the more its proponents compile such evidence, the more the fundamental concepts of Embodied Cognition remain in the dark. This conundrum is nicely exemplified by Pecher and Zwaan’s book, Grounding Cognition, which is a programmatic attempt to rally together an array of empirical results and linguistic data, (...) and its successes in this endeavor nicely epitomize current directions among the various research provinces of Embodied Cognition. The untoward drawback, however, is that such successes are symptomatic of the growing imbalance between experimental progress and theoretical interrogation. In particular, one of the theoretical cornerstones of Embodied Cognition —namely, the very concept of grounding under investigation here—continues to go unilluminated. Hence, the advent of this volume indicates that—now more than ever—the concept of grounding is in dire need of some plain old-fashioned conceptual analysis. In that sense, Embodied Cognition is grounded until further notice. (shrink)

According to Margaret Gilbert, two or more people collectively believe that p if and only if they are jointly committed to believe that p as a body. But the way she construes joint commitment in her account – as a commitment of and by the several parties to “doing something as a body” – encourages the thought that the phenomenon accounted for is not that of genuine belief. I explain why this concern arises and explore a different way of construing (...) joint commitment, in order to avoid the concern. This leads me to propose a revised Gilbertian account of collective belief, according to which two or more people collectively believe that p if and only if they are jointly committed to p as true. (shrink)

Arthur Ripstein’s conception of Kantian freedom has exerted an enormous recent influence on scholars of Kant’s political philosophy; however, the conception seems to me flawed. In this paper, I argue that Ripstein’s conception of Kantian freedom as “your capacity to choose the ends you will use your means to pursue” – your “purposiveness” – is both too narrow and too broad: (1) Wrongful acts such as coercive threats cannot choose my ends for me; instead, such acts wrongfully restrict my perceived (...) options. And (2) rightful changes to the context in which I choose that render my means insufficient for my ends restrict my capacity to choose them. Alternatively, my purposiveness reduces to my entitlements; but then freedom as purposiveness is viciously circular or fails as a new approach to the “devastating” objection that motivates it. (shrink)

This chapter develops an account of epistemic nihilism—roughly, the rejection of truth’s intrinsic or instrumental value in favor of statements that reject or obscure truth to secure an advantage for the speaker—by examining three instances of such nihilism: lying, bullshit, and trolling. It further argues that epistemic nihilism, exacerbated by changes in the media landscape, can pose a significant threat to liberal democratic institutions and ideals by undermining the democratic ideal of good faith engagement on a level playing field, while (...) also encouraging undemocratic actions (e.g., terrorism) among interlocutors who take the nihilist’s claims seriously. Finally, this chapter argues that in extreme cases, we are justified in denying epistemic nihilists a platform from which to speak by drawing a parallel with vexatious litigant laws that deny individuals the right to petition courts for redress on the grounds that abuse of that right results in significant harm to both individuals and the legal system itself. (shrink)

The free-energy principle states that all systems that minimize their free energy resist a tendency to physical disintegration. Originally proposed to account for perception, learning, and action, the free-energy principle has been applied to the evolution, development, morphology, anatomy and function of the brain, and has been called a postulate, an unfalsifiable principle, a natural law, and an imperative. While it might afford a theoretical foundation for understanding the relationship between environment, life, and mind, its epistemic status is unclear. Also (...) unclear is how the free-energy principle relates to prominent theoretical approaches to life science phenomena, such as organicism and mechanism. This paper clarifies both issues, and identifies limits and prospects for the free-energy principle as a first principle in the life sciences. (shrink)