The partnership between a visually impaired runner (VIR) and sighted guide runner (SGR) constitutes a unique sporting dyad. The quality of these partnerships may profoundly impact the sport and physical activity (PA) experiences of visually impaired (VI) people, yet little is known about the experiences of VIRs and SGRs. This study aimed to explore qualitatively the running experiences of VIRs and SGRs. Five VIRs and five SGRs took part in in-depth, semi-structured interviews (M length = 62 minutes) exploring their running (...) journeys and perceptions of running-together. We analysed the dataset using reflexive thematic analysis. Four themes were generated, comprising: becoming and being a running team; a multifaceted intercorporeal experience; running-together promotes change; and disabling social interaction within running. Participants were generally positive about their running experiences, highlighting a range of benefits derived from the activity. Nevertheless, some examples of barriers to participation were also identified. Although the positive experiences described by the runners suggest guided running holds promise to increase PA in VI people, our findings illustrate the importance of directing attention towards developing high-quality relationships between VIRs and guides, alongside reinforcing the need for further change to promote inclusivity. (shrink)

Positions taken in the current debate over free will can be seen as responses to the following conditional: -/- If every action is caused solely by another event and a cause necessitates its effect, then there is no action to which there is an alternative (C). -/- The Libertarian, who believes that alternatives are a requirement of free will, responds by denying the right conjunct of C’s antecedent, maintaining that some actions are caused, either mediately or immediately, by events whose (...) effects could be different, even if they were to recur under identical circumstances. We have here a denial of Laplacian Determinism (LD), according to which the condition of the world at any instant makes only one state possible at any other instant. One prominent defender of this view, Robert Kane, holds that unless an agent’s neural mechanisms operated indeterministicly in forming her character she is not responsible for its manifestations. This requirement is entailed by the principle of “ultimate responsibility” (UR) according to which an act is freely willed only if (a) its agent is personally responsible for its performance in the sense of having caused it to occur by voluntarily doing something that was avoidable and (b) there is no sufficient condition for its performance for which its agent is not personally responsible. The orthodox Compatibilist, adducing counterexamples to UR, argues that even if C and its antecedent are true, free will is possible. Ned Markosian has provided the Compatibilist with yet another response to UR-based objections to her view, wedding it to the notion of agent causation: here the deterministic causing of an act by its agent is necessary and sufficient for its being “morally free.” I criticize this view as inconsistent with the agent causalist's belief that moral responsibility precludes causation of a choice by any power other than the maker thereof: he being its efficient cause. (shrink)

Is cultural evolution needed to explain altruistic selfsacrifice? Some contend that cultural traits (e.g. beliefs, behaviors, and for some “memes”) replicate according to selection processes that have “floated free” from biology. One test case is the example of suicide kamikaze attacks in wartime Japan. Standard biological mechanisms—such as reciprocal altruism and kin selection—might not seem to apply here: The suicide pilots did not act on the expectation that others would reciprocate, and they were supposedly sacrificing themselves for country and emperor, (...) not close relatives. Yet an examination of both the historical record and the demands of evolutionary theory suggest the kamikaze phenomenon does not cry out for explanation in terms of a special non-biological selection process. This weakens the case for cultural evolution, and has interesting implications for our understanding of altruistic self-sacrifice. (shrink)

This inquiry is addressed to two questions: (1) what if any logical relations might exist between the concepts of desire and action (as they and the distinctions to which they commit us are ensconced in ordinary parlance), and (2) what if any causal or significant non-causal (i.e., material) relations might ever exist between instances of desire and action? -/- It is held that any credible move to deal with such questions must initially, and at some length, specify the employment conditions (...) for the terms 'want' and 'desire'. This is accomplished in the first two chapters wherein a set of premises is generated for the argument of the inquiry which is presented in full and concluded in the third and final chapter. Premises generated devolve from treatment of the following major topics: (a) the grammar of 'want' and 'desire', (b) the ontological status of wants, (c) first-person and third-person want ascriptions, (d) the sorts of things which may be desired (desiderabilia), (e) wants and lacks, (f) wants and wishes, (g) the 'conflicts' of desires and reason and desire, (h) four major candidates for logical relations between wants and acts, (i) the issue of causation, and (j) the placement of Desire in an adequate ontology of persons. -/- It is argued that every major candidate for a logical relation between wants and acts (or our warrant for believing that such a connection exists) breaks down upon analysis; that little warrant exists for construing wants as causes of acts; that wants and acts are related in significant non-logical and non-causal (i.e., material) respects; and that any fully adequate theory of human action must undertake ontology, placing the category of Desire squarely in the foreground. This project represents neither a general theory of desire nor a general theory of human action. It is rather conceived as a propaedeutic to any such inquiry. It is essentially revisionist in intent, setting out on the one hand to challenge existing claims about wants and acts and, on the other hand, proposing a more satisfactory approach to the issue. In no sense is it suggested that recent Anglo-American contributions are without merit. It is rather argued that they are often inadequate contributions vis-a-vis the larger requirements for an adequate theory of human action. (shrink)

Charles Murray describes himself as a libertarian, most notably in his short book, What it Means to be a Libertarian. He might more accurately have described himself as having libertarian tendencies. My reading of Simple Justice is that the views it espouses are far more traditionalist than libertarian. Neither traditionalist state-retribution nor modernist state-leniency is libertarian. Nor does either provide as just or efficient a response to crime as does libertarian restitution, including restitutive retribution. Here, I shall respond directly only (...) to Murray's views, rather than also deal with state-leniency. This is because I accept Murray's thesis, without endorsing his specific arguments for it, that state-leniency is disastrous as a response to crimes against persons and their justly acquired property. (shrink)

Das Anliegen der vorliegenden Studie ist der Entwurf eines wirklichkeitstreuen Modells der Kommunikation. Ebenfalls hat uns interessiert, die konstitutiven Regeln der Kommunikation zu bestimmen und einige Wirksamkeitsnormen und moralische Normen, die mit wünschenswerten sozialen Kommunikationsformen assoziierbar sein könnten, zu identifizieren. Die Kommunikation ist ein facettenreiches und zugleich interpretationsoffenes Phänomen, welches zahlreiche unterschiedliche theoretische Modelle erlaubt. Sie kann von den Psychologen als selbstständigen Verhaltenstyp, von den Soziologen als entscheidenden Sozialisierungsfaktor, von den Anthropologen als kulturhervorbringendes und -verbreitendes Mittel, von den Semiologen als (...) Prozess der (Um-)Konfiguration der verschiedenen Bedeutungen von Bedeutungsträgern usw. gedeutet werden. Das vorliegende Interpretationsmodell der Kommunikation ist allgemein, zum Teil auch hermeneutisch, indem es a) den Kommunikationsprozess als solchen und nicht etwa als eine gesonderte Instanz desselben berücksichtigt, b) eine einzige, nämlich die interaktive Perspektive auf die Kommunikation hervorhebt und c) zweckgemäß die Kommunikation in einen normativen, intelligiblen Rahmen überträgt, wobei sich eine empirische Beschreibung der Kommunikationsfakten zwecks Feststellung allgemeingültiger Gesetze von selbst erübrigt. Diesem theoretischen Modell entsprechend ist die Kommunikation eine Interaktion oder eine wechselseitige semiotische Interaktion, innerhalb derer die involvierten Personen mithilfe von Zeichen komplementäre Verhalten manifestieren, die bestimmten (sozialen) Regeln unterworfen sind. Demzufolge ist die Kommunikation kein individuelles Spiel, sondern ein Team-Spiel. Ihre Protagonisten spielen alle genauso wichtige Rollen, und von ihrer Teilnahme hängt beides, Gelingen und Misslingen der Kommunikation, gleichermaßen ab. Nachdem wir den Begriff der Kommunikation im allgemeinen erklärt hatten, galt unser Interesse der Definition und der Klassifizierung der exklusiven Ausführungsmittel des oben beschriebenen Prozesses, das heißt, der Zeichen. Die semiotische Literatur behandelt die Zeichen sowohl als (stoffliche, mentale oder gemischte) Objekte als auch als Funktionen. Für unsere Demarche haben wir es vorgezogen, die Zeichen als in Wirklichkeit existierende Reize zu beschreiben, die auf Grund von sozialen Konventionen stets auf „etwas Anderes” als auf sich selbst hinweisen. Dieses gemeinte „Andere“, die Bedeutung des Zeichens, ist nichts anderes als die Gesamtheit seiner Anwendungen. Die Natur des Verhältnisses von Zeichen und (willkürlicher, notwendiger, durch Analogie oder Ähnlichkeit begründeter) Bedeutung hat uns dann erlaubt, vier Zeichenkategorien auszumachen und zu beschreiben: Wörter, Indizien, Bilder und Sinnbilder. Was die Erläuterung der konstitutiven Faktoren der Kommunikation anbetrifft, haben wir im Einklang mit den Forschungen angesehener Semiotiker mit sechs grundlegenden Parametern gerechnet: Sender, Empfänger, Mitteilung, Code, Kommunikationssituation und Kommunikationskanal. Doch war das wirklich die einzige treffliche Option? Hätte der Kommunikationsprozess nicht ausgehend von anderen – wenigeren oder mehreren – Faktoren analysiert werden können? Gewiss kann die Kommunikation auch mittels weiterer Variablen verstanden werden – und die Fachliteratur ist nicht müde geworden, die Vorzüge anderer Beschreibungsmuster zu zeigen –; von den sechs semiotischen, von Roman Jakobson eingeführten Parametern kann man indes nicht wegsehen. Sie erlauben allemal eine ziemlich genaue Bestimmung der durch die Protragonisten der Kommunikation durchgeführten Handlungen. In Übereinstimmung mit der allgemeinen Methode der Handlungstheorie haben wir jeden konstitutiven Faktor der Kommunikation mit je einem Handlungspaar korreliert, zum einen beim Sender, zum anderen beim Empfänger. Die Berücksichtigung der Sprache und implizite der Kommunikation im Kontext der Handlungstheorie ist keine bahnbrechende Unternehmung, da sie unter anderem schon John Austin, John Searle und Jürgen Habermas als Ausgangspunkt ihrer Analyse der Sprechakte gedient hat. Auch die Korrelation der Kommunikationsparameter mit einer Handlung ist kein Novum, indem sie auf analoge Weise bereits durch die Versuche einiger Semiotiker wie Roman Jakobson und Karl Bühler, die Funktionen der Sprache bzw. des linguistischen Zeichens zu systematisieren, unternommen wurde. Unser origineller Beitrag besteht aus der Korrelation jedes Faktors der Kommunikation mit einer Interaktion, das heißt mit einem Paar von komplementären Handlungen, so dass jeder Handlung des Senders eine Handlung des Empfängers entspricht. In diesem Sinne haben wir Sender, Empfänger, Mitteilung, Code, Kommunikationssituation und Kommunikationskanal in spezifischen Interaktionstypen (Sichentlarven – Nachvollziehen, Einflussnahme – Antwort, Sendung – Empfang, syntaktische Informierung – syntaktisches Verständnis, Bezeichnung – „Mitbezeich-nung“ bzw. Anknüpfung einer soziale Beziehung – Beibehaltung einer soziale Beziehung) inszeniert. Nachdem wir den Kommunikationsprozess als Totalität von sechs Interaktionstypen beschrieben hatten, haben wir auf drei mögliche Ordnungen der Kommunikationsformen verwiesen, wobei die Einteilungskriterien jeweils die verschiedenen gebrauchten Zeichentypen, die Art der Beziehung zwischen Sender und Empfänger bzw. der sich spontan manifestierende soziale Abstand zwischen den Kommunizierenden waren. Indem unsere Demarche darauf abzielte, die Eigentümlichkeiten verbaler vs. nonverbaler, formaler vs. informaler, der individuell, gruppenweise oder öffentlich stattfindenden Kommunikation auszumachen, haben wir die Akzente auf die Pragmatik der Kommunikation gesetzt und somit einige Hinweise auf den Erwerb spezifischer Kommunikationsfähigkeiten geben können. Das von uns vorgeschlagene theoretische Modell hat also nicht nur ein besseres Verständnis der Kommunikation beabsichtigt, sondern auch die Feststellung eines vollständigen Regelwerks der Kommunikation. Diese Regeln wurden als Formen des zu erwartenden sozialen Verhaltens behandelt – sowohl im normativen als auch im antizipativen Sinne – und wurden im Gefolge von John Searle in zwei Hauptklassen eingeteilt: konstitutive und normative Regeln. Die konstitutiven Regeln bestimmen die von ihnen logisch-abhängigen Verhaltensweisen und legen die Bedingungen der Kommunikationsakte fest, während die normativen Regeln die vorgegebenen, unabhängigen Verhaltensweisen berücksichtigen und die wünschenswerten Bedingungen für ihre Vollziehung festlegen. Um die konstitutiven Regeln ausmachen zu können, sind wir davon ausgegangen, dass der Prozess der menschlichen Kommunikation kein naturhafter, sondern ein institutioneller ist. Mit anderen Worten ist die Kommunikation das Ergebnis einer sozialen Vereinbarung und eben kein unabhängiges Phänomen, das nur von außen zu beschreiben wäre. Je nachdem, wie die notwendigen Regeln, welche die soziale Vereinbarung definieren, respektiert oder verletzt werden, haben wir fünf mögliche Situationen identifiziert: die vollkommene Kommunikation, die teilweise geglückte Kommunikation, die Nonkommunikation, die relative und die absolute Antikommunikation, wobei man hinzufügen muss, dass die erste und die letzte Situation Grenzerfahrungen ohne jegliche Referenz darstellen. Die konstitutiven Regeln der Kommunikation wurden mit allen sechs spezifischen Interaktionstypen korreliert (Sichentlarven – Nachvollziehen, Einflussnahme – Antwort, Sendung – Empfang, syntaktische Informierung – syntaktisches Verständnis, Bezeich-nung – „Mitbezeichnung“ bzw. Anknüpfung einer soziale Beziehung – Beibehaltung einer soziale Beziehung), so dass sie wenigstens aus einem bestimmten Gesichtspunkt ein konsistentes und vollständiges System bilden können. Im Unterschied zum System der konstitutiven Regeln sind hier die Kommunikationsnormen nicht erschöpfend behandelt worden. So wurden aus der Vielfalt der Vorschriften, die zu erwünschten Ergebnissen in der Kommunikation führen könnten, nur einige im Rahmen der vorliegenden Studie angegangen. Diese betreffen die Wirksamkeit und die moralischen Aspekte der Kommunikation und wurden ihrerseits mit den oben angeführten sechs Interaktionstypen korreliert. Dabei waren nicht so sehr die unausweichlichen Lücken im Inventarisieren der Normen, welche die Genauigkeit und die Ökonomie der Kommunikation gewähren, am problematischsten, sondern vielmehr der Grad an moralischer Relevanz, den die Kommunikationsakte beanspruchen können. Wir sind fest davon überzeugt, dass die Handlungen der Kommunizierenden eher davon abhängen, inwiefern sie moralische Verpflichtungen eingehen, und eben nicht davon, was für einen Gewinn sie dadurch erzielen, da keine der berücksichtigten Handlungen sich der Moral gänzlich entziehen können. Ob „neutrale Metaregeln“ für den Fall, dass einige moralische Eingrenzungen mit den Wirksamkeitsnormen kollidieren, vonnöten sind, sei dahin gestellt. Ein theoretisches Modell kann nur dann als nützlich bezeichnet werden, wenn es neue Forschungsansätze bringt und neue Forschungswege ermöglicht. Unter diesem Blickwinkel hat die vorliegende Studie zwei mögliche Fortsetzungen: die Ergänzung der Gesamtheit der normativen Regeln der Kommunikation (im allgemeinen), bzw. das Unterteilen des allgemeinen Modells der Kommunikation in stereotypen Mustern wie der politischen oder didaktischen Kommunikation usw. Schließlich hängen Gelingen und Anwendbarkeit der vorliegenden Studie in pragmatischer Sicht eben davon ab, inwiefern sie dem Leser / Benutzer als Wekzeug zur Verbesserung seiner Kommunikationsleistungen dienen kann. (shrink)

Emphasizing the human body in all of its forms, Beauty Unlimited expands the boundaries of what is meant by beauty both geographically and aesthetically. Peg Zeglin Brand and an international group of contributors interrogate the body and the meaning of physical beauty in this multidisciplinary volume. This striking and provocative book explores the history of bodily beautification; the physicality of socially or culturally determined choices of beautification; the interplay of gender, race, class, age, sexuality, and ethnicity within and on the (...) body; and the aesthetic meaning of the concept of beauty in an increasingly globalized world. Foreward by Carolyn Korsmeyer. Authors include Noel Carroll, Gregory Velazco Y Trianosky, Monique Roelofs, Whitney Davis, Eleanor Heartney, Diana Tietjens Meyers, Phoebe M. Farris, Mary Devereaux, Jo Ellen Jacobs, Karina L. Cespedes-Cortes and Paul C. Taylor, Fedwa Malti-Douglas, Stephen Davies, Jane Duran, Valerie Sullivan Fuchs, Keith Lehrer, Allen Douglas, Cynthia Freeland, Eva Kit Wah Man, Mary Bittner Wiseman, and Peg Brand's essay, "ORLAN Revisited: Disembodied Virtual Hybrid Beauty.". (shrink)

This paper aims to suggest a new approach to Plato’s theory of being in Republic V and Sophist based on the notion of difference and the being of a copy. To understand Plato’s ontology in these two dialogues we are going to suggest a theory we call Pollachos Esti; a name we took from Aristotle’s pollachos legetai both to remind the similarities of the two structures and to reach a consistent view of Plato’s ontology. Based on this theory, when Plato (...) says that something both is and is not, he is applying difference on being which is interpreted here as saying, borrowing Aristotle’s terminology, 'is is (esti) in different senses'. I hope this paper can show how Pollachos Esti can bring forth not only a new approach to Plato’s ontology in Sophist and Republic but also a different approach to being in general. -/- Keywords Plato; being; difference; image; pollachos esti; pollachos legetai 1. Being, Not-Being and Difference The three dialogues where the notion of "difference" attaches to the notion of being, namely Parmenides II, Sophist and Timaeus,and specifically the first two we try to discuss here. In these dialogues, Plato is going to achieve a new and revolutionary understanding of being which is not anymore based on the notion of "same" as it was before in Greek ontology. It was his discovery, I think, that the notion of being in the Greek ontology is attached to the notion of the "same" and it is because of this attachment that there have always been many problems understanding being especially after Parmenides. That being has always been relying on the "same" can be found out from the way most of the Presocratics understood it. It was based on such a relationship between being and "same" that a later Ionian, Heraclitus of Ephesus, rejected Being by rejecting its sameness: unable to be the same, being cannot be being anymore but becoming. Heraclitus’ criticism of his predecessors’ understanding of being was due to his discovery that what they call being is not the same but different in every moment. The relation of being and sameness reaches to its highest point in Parmenides. What Plato does in using the "difference" is nothing but the establishment of a creative relation between being and "difference". In this new relation, although he is in agreement with Heraclitus that being is not the same but different, he does not do it by use of becoming. He disagrees, on the other hand, with Parmenides that such a relation between being and difference leads to not being. At Parmenides 142b5-6 it is said that if One is, it is not possible for it to be without partaking (μετέχειν) of being (οὐσίας), which leads to the distinction of being and one: -/- So there would be also the being of the one (ἡ οὐσία του̑ ἑνὸς) which is not the same (ταὐτὸν) as the one. Otherwise, it wouldn’t be its being, nor the one would partake of it. (142b7-c1) -/- The fact that what is (ἔστι) signifies (σημαῖνον) is other (ἀλλο) than what One signifies (c4-5), is being taken as a reason for their distinction. The conclusion is that when we say 'one is', we speak of two different things, one partaking of the other (c5-7). Having repeated these arguments of the otherness of being and one at 143a-b, Parmenides says that the cause of this otherness can be neither Being nor One but "difference": -/- So if being is something different (ἕτερον) and one something different (ἕτερον), it is not by being one that the one is different from being nor by its being being that being is other than one, but they are different from each other (ἕτερα ἀλλήλων) by difference (τῷ ἑτερῳ) and otherness (ἄλλῳ). (143b3-6) -/- The fifth hypothesis, 'one is not' (160b5ff.) is also linked with the notion of difference. When we say about two things, largeness and smallness, that they are not, it is clear that we are talking about not being of different (ἕτερον) things (160c2-4). When it is said that something is not, besides the fact that there must be knowledge of that thing, we can say that it entails also its difference: 'difference in kind pertains to it in addition to knowledge' (160d8). Parmenides explains the reason as such: -/- For someone doesn’t speak of the difference in kind of the others when he says that the one is different from the others, but of that thing’s own difference in kind. (160e1-2) -/- Although the theory of being as "difference" is not fulfilled yet, an exact look at what occurs in Sophist can make us sure that this was the launching step for "difference" to get its deserved role in Plato’s ontology. The notion of the "difference" is not yet well-functioned in Parmenides because we can see that being is still attached to the same: -/- For that which is the same is being (ὄν γὰρ ἐστι τὸ ταὐτόν) (162d2-3). -/- The notion of difference in Sophist is the key element based on which a new understanding of being is presented and the problem of not being is somehow resolved. The friends of Forms, the Stranger says, are those who distinguish between being and becoming (248a7-8) and believe that we deal with the latter with our body and through perception while with the former, the real being (ὄντως οὐσίαν) with our soul and through reasoning (a10-11). Being is then bound with the "same" by adding: -/- You say that being always stays the same and in the same state (ἣν ἀεὶ κατὰ ταὐτὰ ὡσαύτως ἔχειν) but becoming varies from one time to another (δὲ ἄλλοτε ἄλλως). (248a12-13) -/- That the theory of the relation of being and capacity (247d8f., 248c4-5) matches more with becoming than with being (248c7-9) must be rejected because being is also the subject of knowledge which is kind of doing something (248d-e). It does, however, confirm that 'both that which changes and also change have to be admitted as existing things (ὄντα) (249b2-3). I believe that this is what Socrates would incline to do at Theaetetus 180e-181a, that is, putting a fight between two parties of Parmenidean being and Heraclitean becoming and then escaping. The solution is that becoming is itself a kind of being and we ought to accept what changes as being. This is what must be done by a philosopher, namely, to refuse both the claim that 'everything is at rest' and that 'being changes in every way' and beg, like a child, for both and say being (τὸὄν) is both the unchanging and that which changes (249c10-d4). This kind of begging for both is obviously under the attack of contradiction (249e-250b). For both and each of rest and change similarly are (250a11-12) but it cannot be said either that both of them change or both of them rest, being must be considered as a third thing both of the rest and change associate with (250b7-10). The conclusion is that 'being is not both change and rest but different (ἕτερον) from them instead' (c3-4). The notion of difference helps Plato to take being departed from both rest and change because it was their sophisticated relation with being that made the opposition of being and becoming. Plato is now trying to separate being from rest and, thus, from "same" by "difference". Such a crucial change is great enough to need a 'fearless' decision (256d5-6). The possibility of being of not being is resulted (d11-12) comes as the answer to the question 'so it’s clear that change is not being and also is being (ἡ κίνησις ὄντως οὐκ ὄν ἐστι καὶ ὄν) since it partakes in being?' (d8-9). It is then by the notion of difference that becoming is considered as that which both is and is not. This coincidence of being and not being about change is apparently similar to Socrates’ paradoxical statement at Republic 477a about what both is and is not. -/- Introduction The Republic 476-477 has always been a matter of controversy mainly about two interwoven points. The first problem is the meaning of being here; that whether what he has in mind is a veridical, existential or propositional sense of being. The second problem is his distinction between the objects of knowledge and opinion which seems to lead, some believe, to the Two Worlds (TW) theory. The crucial point in Republic is that what is considered between knowledge (ἐπιστήμης) and ignorance (α͗γνοιας), namely opinion, must have a different object that leads Socrates to draw the distinction of knowledge and opinion between their objects. The problem of understanding being in the fifth book of the Republic is that when it is said that the Form of F is F but a particular participating in F, both is and is not F, it sounds too bizarre and unacceptable. It cannot be imaginable how a thing can be existent and non-existent at the same time. At the first sight, the only solution seems to be the degrees of existence which is called by Annas (1981, 197) a 'childish fallacy' and a 'silly argument'. Kirwan (1974, 118) thinks that Republic V does not attribute 'any doctrine about existence' to Plato and Kahn (1966, 250) claims that the most fundamental value of einai when used alone (without predicate) is not "to exist" but "to be so", "to be the case" or "to be true". The problems of understanding being in Republic and Sophist besides the difficulties of the existential reading led scholars to the other senses of being, mostly related to the well-known Aristotelian distinctions between different senses of being. In the predicative reading, Annas, for example, refers this difference to the qualified and unqualified application. Whereas the Form of F is unqualifiedly F, a particular instance of F can be F only qualifiedly (1981, 221). Vlastos’ well-known substitution of 'degrees of reality' for 'degrees of being/existence' must be categorized as a predicative reading. Kahn thinks that the basic sense of being for Plato is 'something like propositional structure, involving both predication and truth claims, together with existence for the subject of predication' (2013, 96). Believing that the complete-incomplete distinction terminology is misleading about Plato, he thinks that semantic functions are only second-order uses of the verb and it is the predicative or incomplete function which is fundamental. Suggesting a veridical reading, Fine (2003, 70 ff) thinks that while both existential and predicative readings separate the objects of knowledge and belief, it is only her reading which does not force such separation of the objects and thus does not imply TW. Stokes (1998, 266) thinks that though Fine is right saying that Plato does not endorse TW in book V, she is wrong in rejecting existential in favor of the veridical reading. The reception of existential reading can be seen more obviously in Calvert who thinks, in agreement with Runciman, that 'it would be safer to say that Plato’s gradational ontology is probably not entirely free from degrees of existence' (1970, 46). At Sophist 254d-e Plato singles out five most important kinds (or Forms!?) in which the same (ταὐτὸν) and difference (θάτερον) are regarded besides being, rest and change. They are, therefore, neither the same nor the difference but share in both (b3). Being (τὸ ὄν) cannot be the same also because if they 'do not signify distinct things' both change and rest will have the same label when we say they are (255b11-c1). We have then four distinct kinds, being, change, rest and same, none of them is the other. The case of difference is more complicated. When the stranger wants to assess the relation of being and difference, he can say simply neither that they are distinct nor that they are not. He has to make an important distinction inside being to get able to draw the relation of being and difference: -/- But I think you'll admit that some of the things that are (τῶν ὄντων) are said (λέγεσθαι) by themselves (αὐτὰ καθ’ αὑτά) but some [are said] always referring to (πρὸς) other things (ἄλλα) (255b12-13) -/- The difference is always said referring to other things (τὸδέγ’ ἕτερον ἀεὶ πρὸς ἕτερον) (255d1). It pervades all kinds because each of them should be different from the others and is so due to the difference and not its own nature (253e3f.) After asserting that change is different from being and therefore both is and is not (256d), the difference is described as what makes all the other kinds not be, by making each different from being. Given that all of them are by being, this association of being and difference is the cause of their being and not-being at the same time, the issue that its version at RepublicV made all those controversies we discussed above: -/- So in the case of change and all the kinds, not being necessarily is (Ἔστιν ἄρα ἐξ ἀναγκης τὸ μὴ ὄν). Τhat’s because as applied to all of them, the nature of the difference (ἡ θατέρον φύσις) makes each of them not be by making it different from being. And we’re going to be right if we say that all of them are not in the same way. And conversely [we’re also going to be right if we say] that they are because they partake in being. (Sophist 256d11-e3) -/- Plato’s new construction of five distinct kinds and the role he gives to thedifference among them is aimed to resolve the old problem of understanding being which has always been annoying from the time of Heraclitus and Parmenides. Both the ontological status of becoming and that of not being were, in Plato’s mind, based on the absolute domination of the notion of the Same over being. Now, not only becoming is understandable as being but also not being which is not the contrary of being anymore but only different (ἕτερον) (257b3-4). Though I agree partly with Frede that the account of not being which is needed for false statements is more complicated than just saying, as Cropsey (1995, 101) says, that Plato is substituting 'X is not Y' with 'X is different from Y', I totally disagree with him that when we say X is not beautiful, Plato could not have thought that it is not a matter of its being different from beautiful because 'it would be different from beauty even if it were beautiful by participation in beauty' (1992, 411). Conversely, as we will discuss, it is exactly the relation of the beautiful thing, X, and the beautiful itself, in which X shares that is to be solved by the notion of not being as difference. Though it is beautiful because of sharing in beauty, X is not beautiful because it is different from beautiful itself. What the difference is to do is to show how something can both be and not be the same thing. The difference is what makes one thing both be and not be a certain other thing. This equips the difference with the ability to explain a certain thing’s not-being when it is. Thanks to the notion of difference, it is now possible to explain not only not being but also the simultaneous being and not being of a thing: 'What we call "not-beautiful" is the thing that ἕτερόν ἐστιν from nothing other than του̑ καλου̑ φύσεως' (257d10-11). The result is that not beautiful happens to be (συμβέβηκεν εἶναι) one single thing among kinds of beings (τι τῶν ὄντων τινὸς ἑνὸς γένους) and at the same time set over against one of the beings (πρός τι τῶν ὄντων αὖ πάλιν ἀντιτεθὲν) (257e2-4) and thus be something that happens to be not beautiful (εἶναί τις συμβαίνει τὸ μὴ καλόν); a being set over against being (ὄντος δὴ πρὸς ὄν ἀντίθεσις) (e6-7). Neither the beautiful is more a being (μα̑λλον ... ἐστι τῶν ὄντων) nor not beautiful less (e9-10) and thus both the contraries similarly are (ὁμοίως εἶναι) (258a1). This conclusion, it is emphasized again (a7-9), owes to θατέρου φύσις now turned out as being. Therefore, each of the many things that are of the nature of the difference and set over each other in being (τῆς τοῦ ὄντος πρὸς ἄλληλα ἀντικειμένων ἀντίθεσις) is being as being itself is being (αὐτοῦ τοῦὄν τοςοὐσία ἐστιν) and not less. They are different from, and not the contrary of, each other (a11-b3). This is exactly τὸμὴὄν, the subject of the inquiry (b6-7). Hence, not being has its own nature (b10) and is one εἶδοςamong the many things that are (b9-c3). Such far departing from Parmenides’ ontological principle is done on the basis of the nature of the difference. It was the discovery of such a notion that made the stranger brave enough to say that not being is each part of the nature of the difference that is set over against being (258d7-e3, cf. 260b7-8). That the relation of being and difference is difference is the key element of the new ontology. The difference is, only because of its sharing in being, but it is not that which it shares in but different from it (259a6-8). Not being is exactly based on this difference: ἕτερον δὲ τοῦ ὄντος ὄν ἔστι σαφέστατα ἐξ ἀνάγκης εἶναι μὴ ὄν (a8-b1). 2. Difference and the Being of a Copy We discussed above that the sense of being of particulars in Republic V made so many debates that whether being is there used in an existential sense or not. Particulars in Republic are regarded as images in the allegories of Line and Cave. The being of an image/copy makes, thus, the same problem. Plato’s analogy of original -copy for the relation of Forms and their particulars in Republic has obviously a different attitude to being. The main question is that what is the ontological status of a copy in respect of its original? Are there two kinds of being, 'real being' versus 'being' as Ketchum says (1980, 140) or only one kind? What is the difference of being in an original and its copy? Is it a matter of degrees of being or reality as some commentators have suggested? Is it a matter of being relational? By reducing the ontological issue to an epistemological one, Vlastos’ suggestion of degrees of reality in an article with the same name does neither, I think, pay attention to the problem nor resolve it. He agrees that Plato never speaks of "degrees" or "grades" of reality (1998, 219). What allows him to entitle it as such are some of Plato’s words in Republic as well as Plato’s words in some other dialogues (1998, 219). When Plato states that the Forms only can completely, purely or perfectly be real he means, Vlastos says, they are cognitively reliable (1998, 229); an obvious reduction of the issue to an epistemological one. He thinks that when in Republic we are being said that a particular’s being F is less pure than its Form, it is because it is not exclusively F, but it is and is not F and this being adulterated by contrary characters is the reason of our confused and uncertain understanding of it (1998, 222). Ketchum rightly criticizes Vlastos’ doctrine in its disparting from ontology thinking that 'to understand Plato’s talk of being as talk of reality is to obscure the close relation that exists between "being" and the verb "to be"' (1980, 213). He thinks, therefore, that οὐσία must be understοοd as being rather than reality, τὸὄν as "that which is" and not "that which is real" and … (ibid). His conclusion is that degrees of reality cannot interpret Plato correctly and we must accept degrees of being. Allen believes that a 'purely epistemic' reading of the passage in Republic is patently at odds with Plato’s text (1961, 325). He thinks that not only degrees of reality but also degrees of reality must be maintained (1998, 67). What Cooper suggests gets close to this paper’s solution: -/- Plato does not I think wish to suggest that existence is a matter of degree in the way in which being pleasant or painful is a matter of degree. Rather there are different grades of ontological status. (1986, 241) -/- A more ontological solution for the problem of understanding the being of a copy and its relation with the being of its original is suggested by the theory of copy as a relational entity. Based on this interpretation, 'the very being of a reflection is relational, wholly dependent upon what is other than itself: the original…' (Allen, 1998, 62). As relational entities, particulars have no independent ontological status; they are purely relational entities which derive their whole character and existence from Forms (ibid, 67). Although these relational entities are and have a kind of existence, we must also say that 'they do not have existence in the way that Forms, things which are fully real, do' (ibid). Allen (1961, 331) extends his theory to Phaedo where it is said that particulars are deficient (74d5-7, 75a2-3, 75b4-8), 'wish' to be like (74d10) or desire to be of its nature (75a2); an extension that, like F.C. white (1977, 200), I cannot admit. He correctly states that Plato did not start out with a doctrine of particulars as images and semblances but come to such a view after Phaedo, or perhaps after Republic V (1977, 202). Though we may not agree with him about Republic V, if we have to consider its last pages also, we must agree with him that not only the ontology of Phaedo but also that of Republic II-V (except the last pages of the latter book) are somehow different from (but at the same time appealing to) the ontology of original-copy which should exclusively assign to Sophist, Timaeus and RepublicVI-VII besides the last pages of book V. The answer to the problem of Plato’s sense of being in RepublicV can be reached only if we read Republic V based on and as following Sophist. We can find out his meaning of that which both is and is not only by the ontological status he assigns to a copy in Sophist. The kind of being of a copy in Sophist reveals as Plato’s key for the lock of the problem of not being. Let’s see how the ontological status of a copy is the critical point of Plato’s ontology. In the earlier pages of Sophist, we are still in the same situation about not being. To think that that which is not is is called a rash assumption (237a3-4) and Parmenides’ principle of the impossibility of being of not being is still at work (a8-9). At 237c1-4, the problem of "not being" is noticed in a new way which shows some kinds of a more realistic position to the problem of not being. Nevertheless, not being is still unthinkable, unsayable, unutterable and unformulable in speech (238c10). Soon after mentioning that it is difficult even to refuse not being (238d), the solution to the problem appears: the being of a copy (εἴδωλον) (239d). A copy is, says Theaetetus, something that is made referring to a true thing (πρὸς τἀληθινὸν) but still is 'another such thing (ἕτεροντοιου̑τον)' (240a8). Nevertheless, this 'another such thing' cannot be another such real or true thing. In answer to the question of the Stranger that if this 'another such thing' is the true thing (240a9), Theaetetus answers: οὐδαμῶς ἀληθινόν γε, ἀλλ’ ἐοικὸςμὲν (240b2). A copy’s being 'another such thing' does not mean another true thing but only a resemblance of it. Not only is not a copy another true thing besides the original, but it is the opposite of the true thing (b5) because only its original is the thing genuinely and being a copy is being the thing only untruly. The word ἐοικὸς is opposed to ὄντως ὄν in the next line (240b7): 'So you are saying that that which is like (ἐοικὸς) is not really that which is (οὐκ ὄντως [οὐκ] ὄν)'. But still a copy 'is in a way (ἔστι γε μήν πως)' (b9). While it is not really what it is its resemblance, it has its own being and reality because it really is a likeness (εἰκων ὄντως) (b11). The Stranger asks: -/- So it is not really what is (οὐκ ὄν ἄρα [οὐκ] ὄντως ἐστὶν) but it really is what we call a likeness (ὄντως ἣν λέγομεν εἰκόνα)? (b12-13) -/- This is Plato’s innovative ontological solution to the problem of not being. Theaetetus’ answer confirms this: 'Maybe that which is not is woven together with that which is' (c1-2). Therefore, a copy neither is what really is nor is not-being but only is what in a way is. Thanks to the ontological status of a copy, the third status intermediate between being and not being is brought forth. The essence of an image, in Kohnke’s words, does not consist 'solely in the negation of what is genuine and has real being' because otherwise 'it would be an ὄντως οὐκ ὄν,essentially and really a not being' (1957, 37). The characteristics of a copy can be summed up as folows: i) A copy is a copy by referring to a true thing (πρὸς τἀληθινὸν). ii) A copy is different from that of which it is a copy (ἕτερον). iii) A copy is not itself a true thing (ἀληθινόν) as that of which it is a copy but only that which is like it (ἐοικὸς). iv) It is not really that which really is (ὄντως ὄν) but only really a likeness (εἰκων ὄντως). The conclusion is that: v) A copy in a way (πως) is that means it both is and in not, the product of interweaving being with not being. This leads to the refutation of father Parmenides’ principle, accepting that 'that which is not somehow is (τό τε μὴ ὄν ὡς ἔστι)' and 'that which is, somehow is not (τό ὄν ὡς οὐκ ἔστι) (241d5-7). Besides copies and likenesses (εἰκόνων), we have also imitations (μιμημάτων) and appearances (φαντασμάτων) as the subjects of this new kind of being and thus false belief (241e3). In Timaeus, the world of becoming which cannot correctly be called and thus we have to call it "what is such" (τὸ τοιου̑τον) (49e5) or "what is altogether such" (τὸ διἀ παντὸς τοιου̑τον) (e6-7), consists solely of imitations (μιμημάτα) (50c5) which are identifiable only by the things that they are their imitations. The word τοιου̑τον which had been used to determine the situation of a copy in respect of its original, now becomes the definition of the world of becoming in which everything is an image of another thing, a Being, that stays always the same and is different and separated from its image. Cherniss, in my view rightly, draws attention to the very important point about the ontological status of an image that can at the same time be considered a criticism of the relational theory. What we are being said in Timaeus, he thinks,cannot be explained by saying that an image is not self-related and making its being relational. What is crucial about an image is that it 'stands for something, refers to something, means something and this meaning the image has not independently as its own but only in reference to something else apart from it' (1998, 296). This function finds its best explanation in the theory we are to suggest in the following. 3. πολλαχῶς ἔστι The best way to understand the ontological status of an image in Plato is to see first how his most clever pupil, Aristotle, resolved the same problem that Plato brought his theory of image for its sake. Aristotle’s theory of pollachos legetai is a brilliant and, at the same time, deviated version of Plato’s theory that is able, however, to help us read Plato in a better way. We discuss Aristotle’s theory to reach to a full understanding of Plato’s theory because it is, firstly, constructed in Aristotle in a more clear way and, secondly, it can also be taken as an evidence that our reading of Plato is legitimate. The phrase τὸ ὄν πολλαχῶς λέγεται, a so much repeated phrase in Aristotle’s works, is his resolution for some of the ontological problems of his predecessors all treating being as if it has only one sense. Aristotle is right in his criticism of the philosophical tradition specially Heraclitus, Parmenides and Plato since all did presuppose only one sense for being and his theory is, thus, a creative and revolutionary solution for many problems that all the past philosophers were stuck in. But it is at the same time somehow a borrowed theory. As we will discuss, both the structure of the doctrine and the problems it tries to resolve are the same as Plato’s doctrine (and even is comparable in its phraseology) though it is in Aristotle, as can be expected, a more clear and better structured doctrine. 1) Associated with the theory of pros hen and the theory of substance, the theory of several senses of being provides a structure which, I insist, is the best guide to understand Plato’s theory of Being in Sophist, Timeaus and Republic. a) Although the theory of pollachos legetai is not necessarily based on the theory of pros hen, they become tightly interdependent about being: -/- Being is said in many ways/senses (τὸ δὲ ὄν λέγεται μὲν πολλαχῶς) but by reference to one (πρὸς ἕν) [way/sense] and one kind of nature (μίαν τινὰ φύσιν). (Metaphysics 1003a33-34) -/- The doctrine of pros hen which is Aristotle’s initiative third alternative besides the homonymous and synonymous application of words, is primarily a linguistic theory that tries to provide a new theory to explain the different implementations of the same word. The pros hen implementation of being is to provide an alternative for the theory of the synonymous (in Plato: homonymous) implementation of being which says being is said in one sense (kath hen) (1060b 32-33). That both the pros hen and the kath hen implementation of a word has one thing (hen) as what is common, makes them in opposition to the homonymous implementation which does not consider anything in common. Whereas both pros hen and kath hen assume a common nature, with which all the implementations of the word have some kind of relation, their difference is that while kath hen takes all the implementations of the word as the same with the common nature, pros hen makes them different. Substance is called πρῶτον ὄν because it is said to be primarily: -/- For as is (τὸ ἔστιν) is predicated of all things, not however in the same way (οὐχ ὁμοίως) but of one sort of thing primarily and of others in a secondary way. So too τὸτί ἐστιν belongs simply (ἁπλῶς) to substance but in a limited sense (πῶς) to the others [other categories] (1030a21-23). -/- The word ἁπλῶς standing against κατὰ συμβεβηκός tries to make substance different from the accidents. When we are being said that τὸ ὄν πολλαχῶς λέγεται, it means that only the substance that is simply (ἁπλῶς) the ἕν, the common nature, τὸὄν. When we use the word 'being' about a substance, the being is said differently from when we use 'being' about an accident. The distinction between the substance and the other categories is a distinction between what simply is said to be and what only with reference to (pros) the substance is said to be. The doctrine of pros hen, changing kath hen to pros hen in respect of to on, makes a distinction that wants to show that while there is a kind of implementing the word being that is simply being, there is another kind which is called being only by reference to that which is simply being. In the doctrine of pros hen it is not so that all the things that are said to be are only by reference to a common one thing, but that while one thing is called being because it is that thing itself, the other things are called so without being that thing itself but only by referring to it. At the very beginning of book Γ, it is said that: -/- Being is said in many senses but all refer to one arche. Some things are said to be because they are substances, others because they are affections of substances, others because they are a process towards substances or destructions or privations or qualities of substances … (1003b5-9, cf. 1028a18-20) -/- Substance is what really is said to be and all other things that are said to be are said only in favor of it. This difference of substance from all other senses of being is what is, I believe, primarily aimed in Aristotle’s interrelated theories of pollachos legetai,pros hen and the theory of substance. b) The difference of the implementation of being in the case of substance and the accidents goes so deep that while substance is considered as the real being, the accidents are almost not being. An accident is a mere name (Metaphysics 1026b13-14) and is obviously akin to not being (b21). Aristotle adds that Plato was 'in a sense not wrong' saying that sophists deal with not being (τὸ μὴ ὄν) because the arguments of sophists are, above all, about the accidental (1026b13-16). At the beginning of book , he says about quality and quantity (which look to be more of a being than other accidents) that they are not existent (οὐδ̕ ὄντα ὡς εἰπεῖν) in an unqualified sense (ἁπλῶς) (1069a21-22). The two above-mentioned points, Aristotle’s (a) interwoven theories of pollachos legetai, pros hen and the theory of substance and (b) taking accidents almost as not being, comparedwith substance, brings forth a structure that I shall call Pollachos Legetai (with capital first letters). What is of the highest importance in this structure for me is the difference of substance from accidents and the kind of relation which is settled between them. There is a substance that without any qualification is said to be and the accidents that are said to be only by reference (pros) to it. Adding Aristotle’s point about accidents that they are nearly not being to this relation and difference, we can obviously see how much this structure is close to Plato’s original-copy ontology. We spoke of the relation of being and difference in Plato’s model and the way Plato construes the being of a copy. A copy is a copy only by referring to (pros) a model; it is different from (ἕτερον) that of which it is a copy; it is not itself a true thing as its model and not really that which is (ὄντως ὄν) but only is in a way (πῶς). If we behold the difference of substance and accident in the context of the theory of pollachos legetai and pros hen, we can observe its fundamental similarity with Plato’s original-copy theory in its structure. Allen draws attention to the fact that the relation between Forms and particulars in Plato’s original-copy model is 'something intermediate between univocity and full equivocity' (1998, 70, n. 24) and the same as what Aristotle calls it pros hen (ibid). What made us compare the two structures was not, of course, the complete similarity of two structures (we have to agree with many possible differences of the two theories) but exactly the specific relation between an original and its copy on the one hand, and a substance and its accident on the other hand. As substance and accident do not share a common character and the substance -accident model hints that they stand in a certain relation, there is no common character between the original and copy in Plato’s model as well. Furthermore, their similarity is not confined to their structure only; they are also aimed to solve the same problem. The central point of the theory is that all the predecessors took being in one sense and this was their weakness point. Besides the mentioned above passages about the relation of pollachos legetai and presocratics’, as well as Plato’s, ontology, the relation of the theory with the problem of not being is clear in several passages. In Metaphysics, it is said: 'Being is then said in many senses… It is for this reason that we say even of not being that it is not being' (1003b5-10). Discussing the accidental sense of being, Aristotle points that it is in the accidental way that we say, for example, that not-white is because that of which it is an accident is (1017a18-19, cf. 1069a22-24). We mentioned that he thought Plato was right saying that sophistic deals with not being because sophistic deals with accidental, which is somehow not being (1026b14-16). Plato turned sophistic not-being to what both is and is not and Aristotle to what accidentally is said to be. What helps Aristotle to resolve the problem of not being is his distinction between ἁπλῶς and κατὰ σθμβεβηκός. Aristotle’s "qua" (ᾕ) which is directly linked with his distinction between καθ’ αὑτο λέγεται and κατὰ συμβεβηκός λέγεται, is used to resolve the old problem of coming to be out of not being (Physics 191b4-10). He strictly asserts that his predecessors could not solve the problem because they failed to observe the distinction of "qua itself" from "qua another thing" (b10-13). He then continues: -/- We ourselves are in agreement with them in holding that nothing can be said simply (ἁπλως) to come from not being (μὴὄντος). But nevertheless we maintain that a thing may come to be from not being in an accidental way (κατὰ συμβεβηκός). For from privation which ὅ ἐστι καθ’ αὑτο μὴ ὄν, nothing can become. (Phy. 191b13-16, cf. b19-25) . (shrink)

Predication is a lingual relation. We have this relation when a term is said (λέγεται) of another term. This simple definition, however, is not Aristotle’s own definition. In fact, he does not define predication but attaches his almost in a new field used word κατηγορεῖσθαι to λέγεται. In a predication, something is said of another thing, or, more simply, we have ‘something of something’ (ἓν καθ᾿ ἑνὸς). (PsA. , A, 22, 83b17-18) Therefore, a relation in which two terms are posited (...) to each other in a way that one is said (predicated) of the other is a predication. The term of which the other term is said is called a subject (ὑποκειμένον) and the other, which is said about the subject, is called the predicate. Thus, in a predication the predicate is predicated of the subject; given that being predicated is almost as the same as being said. The relation between being said and being predicated is so close that ‘if something is said of a subject both its name and its definition are necessarily predicated of the subject.’ (Cat., 5, 2a19-26) This, however, is true only about the second genera and not the accidents. a) Nature of relation in predication What is Aristotle’s theory about the nature of the relation in a predication? How does he fundamentally understand this relation? Phil Corkum distinguishes between predication logic and traditional term logic and argues that the relation between subject and predicate in Aristotle is of the latter kind. While in predicate logic, subjects and predicates have distinct roles, they have the same role in traditional term logic. In predication logic, subjects refer, but predicates characterize. Thereupon, a sentence expresses a truth if the object to which the subject refers is correctly characterized by the predicate. In traditional term logic, both subjects and predicates refer and a sentences expresses a truth if both name one and the same thing. He concludes that Aristotle ‘problematically conflates prediction and identity claims’ because while he thinks both subjects and predicates refer, he would deny that a sentence is true just in case the subject and the predicate name one and the same thing. Based on this, Corkum believes that Aristotle’s core semantic is not identity but the weaker relation of constitution, which is a mereological interpretation: ‘All men are mortal’ is true just in case the mereological sum of humans is part of the mereological sum of mortals. b) Aristotle’s theory of predication: one or two theories? Frank Lewis finds an inconsistent gap between the theory of predication in the Categories and that in the later books of the Metaphysics VII, 6. So too Joan Kung, Terry Irwin, Daniel Graham. c) Distinction of ‘being said of’ and ‘being in’ Owen enumerates the texts in which Aristotle’s distinction between ‘being said of’ and ‘being in’ is asserted: OI. 11b38-12a17; To., 127b1-4; Cat. 1a20-b9; 2a11-14; 2a27-b6; 2b15-17; 3a7-32; 9b22-24 d) Predication in Aristotle and the standard ‘S is P’ Marie De Rijk Lambertus thinks that the ‘S is P’ pattern is misleading when it comes to express predication in Aristotle. In his view, ‘The Aristotelian procedure should be described in terms of appositively assigning an attribute (κατηγορούμενον) to a substrate (ὑποκείμενον), rather than ascribing a predicate to a subject by means of a copula…. The comment should be considered an attribute which is said to fall to a substrate, without understanding this procedure in terms of sentence predication.’ e) Aristotle’s predication: bipartite or tripartite? It is a difficulty to make Aristotle’s theories of name-verb predication and tripartite predication consistent. The reason is that tripartite is not consistant with his name-verb structure based on which he says that the predicate term is the ‘verb.’ (20a31; 20b1-2; 16a13-5) The problem is that in a tripartite form like ‘Socrates is white’ we cannot take ‘is white’ as the verb because, based on Aristotle himself, no part of a verb can be significant itself. (16b6-7) However, his assertions about the equivalences between the two structures (OI. 12, 21b9-10; PrA. 51b13-16; Met., 1017b22-30) must mean that they are not inconsistent in his own view. Thus, as Allen Bāck points, ‘Aristotle seems to think that he has a single, consistent theory.’ The sense of saying or speaking as the root sense of rhema makes the relation between predication (saying something of something) and verb more interesting. The use of rhema in the sense of a long expression approves this. In Plato’s work (399ac) where Socrates claims that the name anthropos derives from anthron ha opopen (‘he who examines what he has seen’) we have an onoma from, or in place of, a rhema. 1) Subject The subject (ὑποκειμένον) is that of which another term is said or predicated. Aristotle’s own definition is of a much more philosophical value: ‘By subject I mean that which is expressed by an affirmative term (λέγω δὲ ὑποκείμενον τὸ κάταφάσει δηλούμενον).’ (Met., K, 1067b18) Throughout Aristotle’s work, three kinds of subjects can be found: possible, prime and absolute subject. a) Possible subject Those things that can take the position of a subject in a predication we call possible subjects, no matter they can or cannot take the position of predicate in any predication. All terms except the ultimate predicates are possible subjects because they can take the position of subject. b) Prime subject Those that in any predication can only take the position of a subject and not the position of a predicate are prime subjects. (Cat., 5, 3a36-b2; Met., Z, 1028b36-1029a1) This is the truest sense of subject and belongs only to substances. (Met., Z, 1029a1-2) In fact, ‘that which is not predicated of a subject, but of which all else is predicated’ is a substance. (Met., Z, 1029a7-9) In Categories, besides primary substances (Cat., 2, 1b3-6), individual qualities are also said not to be able to be said of anything else. (Cat., 2, 1a23-29) Aristotle’s examples are a certain ‘knowledge-of-grammar’ (τὶς γράμματικὴ) and ‘an individual white’ (τὸ τὶ λευκὸν). Thus, although substances are indeed in the truest sense individuals and, thereby, in the truest sense primary subjects, other individuals can take the position of subject as well. c) Absolute subject While substances are prime subjects of all other things, there is still something of which substances can be predicated and this predication is not an accidental predication: matter. (Met., Z, 1029a21-24) This predication is, indeed, the predication of a ‘form or this’ on matter and material substance. (Met., Θ, 1049a34-36) The only thing that is absolutely a subject, therefore, and can be a predicate is prime matter (πρώτη ὓλη). (Met., Θ, 1049a24-27) d) Subject and substance The most important thing in being a substance is being a subject: ‘It is because the primary substances are subjects for all the other things and all the other things are predicated of them or are in them, that they are called substances most of all.’ (Cat., 5, 2b15-17) And what is nearer to this position is more a substance as a species is more a substance than a genus.’ (Cat., 5, 2b7-8) e) Primary versus accidental subject A subject is a primary subject in a predication when the predicate is predicated of it as itself. In such a case, the subject is the subject of the predicate qua itself and not qua something else. A subject, on the one hand, is an accidental subject when it is not the primary subject of the predicate that is predicated on it. It means that a subject is an accidental subject when the predicate is predicated of it not qua itself but qua something else. An accidental subject is, therefore, anything other than a substance. 2) Predicate as universal It is evident from our discussion of kinds of subjects that except matter, primary substances (if we ignore both accidental predication and cases of absolute subjects) and other individuals, everything else can take the position of a predicate. Since primary substances are individuals, it results that everything except matter and individuals are capable to take the position of predicate. The immediate consequence of this is that the predicate must be a universal. When individuals cannot take the position of predicate, this position cannot be taken by any particular. Therefore, only universals can be predicated of others. Moreover, albeit universals can be a simple subject (for another universal), they are not prime subjects. The closeness of universal and predicate is to the extent that Aristotle differentiates universal from subject. (Met., Θ, 1049a27-30) 3) Categories or classes of predicates Aristotle distinguishes ten highest classes into one of which each predicate must necessarily fall: substance (or what a thing is), quality, quantity, relation, place, time, position, state, activity and passivity. (Cat., 4, 1b25-27; To., I, 9, 103b20- ; PsA., A, 22, 83b13-23) The substance counted among predicates must refer to secondary and not primary substances not only because Aristotle insists that primary substances cannot be predicated but also from his own examples of the category of substance: man and animal. (To., I, 9, 103b25) However, if a predicate be asserted of itself or its genus be asserted of it, the predicate signifies substance or what is; but if ‘one kind of predicate is asserted of another kind,’ it signifies one of the other nines. (To., I, 9, 103b30) These categories are so comprehensive that no predicate can remain outside them so that even non-being has as many senses as categories. (Met., M, 1089a26-30) a) That whether Aristotle’s categories are merely linguistic or fundamentally ontological is a so much controversial dispute. Some like G. E. R. Lloyd believe that ‘categories are primarily intended as a classification of reality … rather than of the signifying terms themselves.’ b) Although Aristotle’s categories have been historically regarded as a classification of predications, there are some recent commentators like Jonathan Barnes that think it is classifying predicates and not predications. 4) Kinds of predicates In his introduction of Categories, as J. W. Thorp truly points out , Aristotle distinguishes between the category of substance and the other categories using μὲν ... δὲ construction. This construction illustrates that beside tenfold classification of categories, there is an even more crucial differentiation between two kinds of predicates: substance or what is on the one hand and the other nine ones on the other hand: the distinction between what is predicated of the subject as what it is and what is predicated of it as an accident. Therefore, we have three kinds of classification - a twofold, a fourfold and a tenfold-complicatedly classifying the same thing, viz. the predicate. Predicates can be divided also to four kinds: property, definition, genus and accident. (To., I, 4, 101b11-20) A property is that predicate that is convertible with its subject and does not signify its essence. A definition is that predicate that is convertible with its subject and signifies its essence. It is a genus if it is not predicated convertibly and is contained in the definition of its subject. And it is an accident if it is neither convertible nor contained in the definition of its subject. (To., I, 8, ^103b3) All of these four kinds are among categories. (To., I, 9, 103b20) It seems that there might be some kind of relation between the twofold and the fourfold distinction: whereas genus and definition are predicated as what it is, accidents are not so predicated. There seems to be some rules that determine to each of these four kinds each predicate belongs. At To., IV, 123b30ff. Aristotle asserts: ‘If B has a contrary and A does not, then B does not belong to A as its genus.’ There is a dispute around per se accidents (τὰ καθ’ αὑτὰ συμβεβηκότα) (Topics., 102a18) whether they must be regarded as either of the four kinds or as a fifth kind. Barnes (1970) argued that they ‘do not fit at all neatly into the fourfold classification.’ He argues that based on the definition of accidents at A, 5, 102b4-5, they must be accidents but based on another definition, A, 5, 102b6-7, they cannot be accidents. He also defends the view that they are not properties. Barnes concludes: ‘This shows that the two definitions are not equivalent, and hence that the ‘predicables’ are not well-defined.’ Demetris J. Hadgopoulos defends the view that the two definitions of accidents are equivalent and per se accidents are properties. 5) Five types of predications We have five kinds of predications based on our division of subjects and our discussion of predicates: simple, primary (itself divided to substantial and accident), accidental (itself divided to primary and secondary), aoincidental and absolute predication. a) Simple predication. This is a predication in which a universal is predicated of either a particular or a universal subject. This sense of predication includes all other senses except the first kind of accidental predication. b) Primary predication. This is a predication in which a universal is predicated of a primary subject, namely a substance. A primary predication is of two kinds: i) Substantial predication. A primary predication in which the predicate is part of, or is included in, the definition of the subject. The universal that is the predicate here is the definition or the genus or the species or the diferentia of the subject. It is this predication that Aristotle calls a unity: ‘A statement may be called a unity … because it exhibits a single predicate as inhering not accidentally in a single subject.’ (PsA., B, 10, 93b35-37) It is only substantial predicates that can be predicated of each other: ‘Predicates which are not substantial are not predicated of one another.’ (PsA., A, 22, 83b18-19) Aristotle defines a substantial predication also in another way. A predication in which a higher genus is predicated on a lower genus, species, a substance or an individual, (or a species is predicated on an individual) is a substantial predication. Therefore, a substantial predication is a predication in a series that has individuals on one of its ends and a general category on its other end. In fact, only substantial predicates can be predicated of one another. (PsA., A, 22, 83b17-19) This series cannot be an infinite series because otherwise not only substances would not be definable but also a genus would be equal to one of its own species. (PsA., A, 22, 83b7-10) -/- ii) Accident predication. A primary predication in which the predicate is not part of, or is not included in, the definition of its subject. The universal that is the predicate here is the property or the accident of the subject. c) Accidental predication. This is a predication in which the subject is not a primary subject and its predicate is a substance. This predication is of two kinds: i) Primary accidental predication. This is a predication in which an accident is predicated of substance. Such a predication can go ad infinitum, which is inferable from Aristotle’s assertion that the secondary accidental predication cannot go ad infinitum. ii) Secondary accidental predication. It is a predication in which an accident is predicated of another accident. Such a predication, Aristotle asserts, cannot go ad infinitum because even more than two accidents cannot be combined. Now what is the meaning of Aristotle’s assertion that we cannot continue this ad infinitum? Does it mean that we cannot continue the predication ‘The musician is white’ and say ‘The white is Athenian?’ But why not? Or maybe he means that by adding any predication, we are still in a condition of predicating two accidents of a substance on each other. -/- As primary subjects, substances can also take the position of predicate though not essentially but accidentally. In propositions like ‘the white is a log’ or ‘That big thing is a log’ we observe substance taking the position of predicate but this is only accidentally: ‘When I affirm ‘the white is a log,’ I mean that something which happens to be white is a log- not that white is the subject in which log inheres (οὐχ ὡς τὸ ὑποκείμενον τῷ ξύλῳ τὸ λευκόν ἐστι), for it was not qua white or qua a species of white that the white (thing) came to be a log (οὔτε λευκὸν ὄν οὔθ᾿ ὃπερ λευκόν τι ἐγἐνετο ξύλον), and the white (thing) is consequently not a log except incidentally.’ (PsA., 22, 83a3-9) Aristotle compares this accidental use of substance in the position of predicate with the substantial use of it in the place of a subject: ‘On the other hand, when I affirm ‘the log is white,’ I do not mean that something else, which happens to be a log, is white (οὐχ ὃτι ἓτερόν τί ἐστι λευκόν, ἐκείνῳ δὲ συμβέβηκε ξύλῳ εἶναι), (as I should if I said ‘the musician is white,’ which would mean ‘The man who happens also to be a musician is white’); on the contrary, log is here the subject- the subject which actually came to be white, and did so qua wood or qua a species of wood and qua nothing else.’ (PsA., 22, 83a8-14) Aristotle asks us not to call propositions like ‘The white is a log’ a predication at all or at least call them accidental predication instead of simple (ἁπλως) predication. (PsA., A, 22, 83a14-17) An accidental predication, in which we have a substance in the place of predicate, is different from an essential predication in that while the subject of an accidental predication is said to be the predicate not by itself but as something different with which it coincides, the subject of an essential predicate is said to be the predicate by itself and not because it is something else: ‘Since there are attributes which are predicated of a subject essentially and not accidentally- not, that is, in the sense in which we say ‘That white (thing) is a man,’ which is not the same mode of predication as when we say ‘The man is white’: the man is white not because he is something else but because he is man, but the white is man because ‘being white’ coincides with ‘humanity’ within one subject. Therefore, there are terms such as are essentially subjects of predicates. (PsA., A, 19, 81a24-29) This capability of ‘non-essentially and only accidentally’ being a subject does belong, in fact, to all sensible things. (PsA., A, 27a32-36) d) Coincidental predication. this is a predication in which none of the subject and predicate are a substance or an individual. In this predication, one of the accidents of a substance or individual is predicated of one of the other accidentals of the same substance or individual. For example, when it is said that ‘The white is musical,’ there is an individual, say Socrates, for which both of white and musical are accidents. e) Absolute predication. This is a predication in which a form or a substance is predicated of a matter or a material substance. Some commentators like Loux and Lewis regarded this predication as close to accident predication, both based on inherence. As R.M. Dancy points out, these predications make Aristotle’s theory of predication immanentist: not only accident predications are explained by the immanence of accidents in substances, some of the substantial predications, which were not explained in Categories, are also explained by the immanence of form in matter. It is interesting that in this kind of predication, it is the predicate and not the subject, which is in the strictest sense substance. As the central books of Metaphysics claim, the form is the substance. Thus, in this predication, substance gets away from the sense of ὑποκείμενον. Corkum mentiones 68a19 as the only instance where Aristotle explicitly claims that a term may be predicated of itself, a passage that is, in Corkum’s view, problematic. 6) Demonstrable predicates Those predicates are demonstrable that are so related to their subjects that there are other predicates prior to them predicable of their subjects (ἔτι δ᾿ ἄλλος, εἰ ὧν πρότερα ἄττα κατηγορεῖται). (PsA., A, 22, 83b32-34) 7) Series of predications Since it is possible for a predicate to be itself the subject of another predication, we can have a series of predications in which the predicate of a predication is the subject of the next predication. This series has the following features: a) It has a limit (in number) (PsA., A, 22, 83b14-16) on the side of subjects: there are subjects that cannot themselves be predicated (PsA., A, 27, 43a39-41), which are, as we noted, prime subjects: particulars, i.e. prime substances and other individuals. (PsA., A, 19-22) Aristotle calls them ultimate (ὓστατον). (PsA., 21, 82a36-b1) b) It has a limit (in number of kinds) (PsA., A, 22, 83b14-16) also on the side of predicates: there must be predicates that cannot be subjects. (PsA., A, 19-22; PsA., A, 27, 43a36-39) Aristotle calls them primary (πρῶτον). (PsA., A, 21, 82b1-4) These are the highest categories or the highest genera of categories. (PsA., A, 22, 83b14-16) c) The series has an escalating shape from mere subjects at its downside to mere predicates at its upside. It is Aristotle himself whos uses the words up and down in this sense. (e.g. PsA., A, 22, 83b2-3) d) The predications that lie between lowest and highest ones must be finite in number. (PsA., A, 19-22 especially: 20, 82a21-35) These have subjects and predicates, each capable of both of the roles of being subject and being predicated. A result of this fitness is that neither demonstration can go to infinity nor everything is demonstrable, the two points Aristotle always insist on. e) The upward side includes the more universal ones and the downward the more particular ones. (PsA., 20, 82a21-23) f) It follows from the above features that ‘neither the ascending nor the descending series of predications … are infinite.’ (PsA., A, 22, 83b24-25) g) Reciprocation and convertibility. Except in case of terms (i.e. subjects and predicates) that are at each of the ends of series of predications, namely ultimates and primaries, it is possible to reciprocate (ἀντιστρέφειν) terms and convert the predication. (PsA., A, 19, 82a15-20) h) Antipredication. Antipredication means that in a predication (S is P), the subject becomes the predicate of its predicate (P is S) in the same category its predicate was predicated on it. For example, if P is in the category of quality, antipredication means that S be predicated of P in the category of quality. In other words, P is a quality of S and S is a quality of P. Aristotle rejects this. (PsA., A, 22, 83a36-b3) i) Self-predication versus other-predication. Aristotle distinguishes between a predication in which a term is said of itself and a predication in which a term is said of another. (Phy., Δ, 2, 209a31-33) j) Predicablity (Cat., 5, 3a36-b2): 1) Primary substances and individuals are not predicable. 2) Secondary substances are of two kinds: genus and species i) Genus is predicable able both of the species and of the individuals. ii) Species is predicable of the individual. 3) Differentiae are predicable both of the species and of the individuals. 8) Predication as classification a) Richard Patterson believes that Aristotle’s so repeated construction using hoper (estin A hoper B) in Topics (120a23 sq., 122b19, 26sq., 123a, 124a18, 125a29, 126a21, 128a35. Also in Posterior Analytics (83a24-30) (Brunschwig’s list. ‘expresses the fact that A is a kind of B (esti A B tis), that A is a species of the genus B.’ 9) Universal predication Aristotle defines universal predication (κατὰ παντὸς κατηγορεῖσθαι) as such: ‘wherever no instance of the subject (τῶν τοῦ ὑποκειμέμνου) can be found of which the other term cannot be asserted.’ (PrA., A, 24b27-29) In this predication, the subject is included in another as in a whole (ἐν ὃλῳ εἶναι) and the predicate is predicated of all of the subject (κατὰ παντὸς κατηγορεῖσθαι). (PrA., A, 24b26-27) Attach another discussion we had about ἐν ὃλῳ here. 10) Quantity of predication A predication, truly stated, has a quantity, which is the number of objects under the name of the subject of which the predicate is predicated. The quantity of subject can be stated in four ways: a. Indefinite: when the predicate belongs or does not belong to the subject without any mark to show to haw many of the particulars under the name of the subject it does or does not belong. E.g. ‘Man is white’; ‘Man is not white.’ (PrA., A, 24a20; OI., I, 7, 17b8-12) b. Universal quantity: When the predicate belongs to all or none of the subject. E.g. ‘Every man is animal’; ‘No man is animal.’ (PrA., A, 24a18-19; OI., I, 7, 17b5-6) The contrary of a predication of a universal quality is a predication of a universal quality. E.g. the contrary of ‘Every man is white’ is ‘No man is white.’ (OI., I, 7, 17b20-23) c. Particular quantity: When the predicate belongs (or does not belong) to some of the subject. E.g. ‘Some men are white’; ‘Some men are not white.’ (PrA., A, 24a19-20) d. Single quantity: when the subject is a proper name of only one object. E.g. ‘Socrates is white’; ‘Socrates is not white.’ The contrary of this predication is of a single quantity: ‘Socrates is not white.’ (OI., I, 7, 17b38-18a4) 11) Convertibility of predication. Some predications are convertible, that is, it is possible to change the place of subject and predicate in a true proposition so that the converted proposition remains true. This is supposed to mean that given the truth of a predication, the truth of the converted predication is inferable. The convert form of a predication depends on its quantity. a. Indefinite quantity: This predication has no strictly true convert because its quantity is not stated. b. Universal quantity: It can be converted in two ways: i) A negative universal can be converted to a negative universal in which the terms are changed. E.g. ‘No pleasure is good’ can be converted to ‘No good is pleasure.’ (PrA., A, 2, 25a5-8 and a14-19) ii) An affirmative universal can be converted to an affirmative particular in which the terms are changed. E.g. ‘Every pleasure is good’ can be converted to ‘Some good is pleasure.’ (PrA., A, 2, 25a7-9) c. Particular quantity: If it is negative, it cannot be converted but if it is affirmative, it can be converted to an affirmative predication with particular quantity. E.g. ‘Some pleasure is good’ can be converted to ‘Some good is pleasure.’ (PrA., A, 2, 25a10-13 and a20-24) d. Single quantity: It cannot be converted. 12) Characteristics of relations between subject and predicate 1. A predicate is of a wider range than its subject. It is based on this fact that Aristotle: a) Prevents individuals to be predicate because there is nothing of which an individual be of a wider range. In other words, it is due to the fact that since an individual is only ‘one particular’ thing and, thus, cannot be of a wider extent than anything that Aristotle prevents them of being a predicate. Matthews, however, thinks we cannot use Socrates, a substance and an individual, in the place of predicate and say it of Socrates because ‘Socrates does not classify Socrates: it names him.’ b) Prevents differentia, species and things under species to be predicated of genus. (To., Z, 6, 144a27-) c) Prevents the species and the things under it to be predicated of the differentia. (To., Z, 6, 144b1-4) d) Also about the effect because it is wider than its subject. (PsA., B, 17, 99a) e) Each attribute is wider than every individual it is predicated on, though several attributes, collectively considered, might not be wider but exactly the substance of a thing. (PsA., B, 13, 96a32-b1) 2. The predicate of a predicate of a subject will be predicated of the subject too: ‘whenever one thing is predicated of another as of a subject, all things said of what is predicated will be said of the subject too.’ (Cat., 3, 1b10-15) In fact, it is due to its predication of the subject that it is predicated of its predicate. (Cat., 5, 2a36-b1) Moreover, what is not predicated of the predicate of a subject cannot be predicated of it as well. (PrA., A, 27, 43b22-27) Thus, what, for example, is not predicated of animal, cannot be predicated of man. 3. The predicate of a subject can be predicated of its predicates as well. (Cat., 5, 3a1-6) For example, you call the individual man grammatical and, thence, you call both a man and an animal grammatical. Nonetheless, the predicate of a subject belongs to it more properly than to its higher predicates. (PrA., A, 27, 43a27-32) 4. It is only the subject that can be distributed and not the predicate. (PrA., A, 27, 43b16-22; OI, I, 7, 17b12-16) Therefore, we can say e.g. ‘Every man is animal’ but we cannot say ‘Every man is every animal.’ 5. It is the reason of the relation between subjects and predicates, that is the reason of predication, which is the subject of inquiry. (Met., Z, 1041a20-24) In other words, since it is a meaningless inquiry to ask why a thing is itself (Met., Z, 1041a14-15), the only remaining meaningful inquiry is to ask why something is something else, i.e. to ask about the reason of predication. 6. A subject cannot categorize its predicate in the same category in which it is categorized by it. If e.g. A is a quality of B, B cannot be a quality of A. Therefore, there is no reciprocation in the same category. (PsA., A, 22, 83a36-39) 13) Characteristics of series of predications 1. A series of secondary accidental predication cannot go ad infinitum for not even more than two terms can be comnbined. For an accident is not an accident of an accident, unless it be because both are accidents of the same subject. (Met., Γ, 1007b1-4) 2. Infinite series cannot be traversed in thought. (PsA., A, 22, 83b6-7) 3. The predications of genera on each other must be ended and cannot go to infinity because otherwise not only substances would not be definable but also a genus would be equal to one of its own species. (PsA., A, 22, 83b7-10) Therefore, a series of predication of genera on each other must be limited on both sides. There must be an upward limit in general categories as well as a downward limit in individual because they cannot be predicated of others. Whatever lies between these limits can both be predicated of others and others be predicated of them. (PrA., A, 27, 43a36-43) 4. The order of predicates matters: it makes a difference whether the series be ABC or BAC. (PsA., B, 13, 96b25-32) . (shrink)

GAMETOGÊNESE -/- Emanuel Isaque Cordeiro da Silva Instituto Agronômico de Pernambuco Departamento de Zootecnia – UFRPE Embrapa Semiárido -/- • _____OBJETIVO -/- Os estudantes bem informados, estão a buscando conhecimento a todo momento. O estudante de Veterinária e Zootecnia, sabe que a Reprodução é uma área de primordial importância para sua carreira. Logo, o conhecimento da mesma torna-se indispensável. No primeiro trabalho da série fisiologia reprodutiva dos animais domésticos, foi abordado de forma clara, didática e objetiva os mecanismos de diferenciação (...) sexual dos embriões em desenvolvimento, quais os genes envolvidos nesse processo e tudo mais. Nesse segundo trabalho, a abordagem será teórica, mas também clara, sobre a formação primordial dos gametas femininos e masculinos, através da ovogênese nas fêmeas e a espermatogênese nos machos. Esse trabalho visa levar a importância do processo de formação dos gametas e a produção hormonal das gônadas, bem como o entendimento sobre as interações com o eixo hipotálamo-hipofisário. -/- •____INTRODUÇÃO -/- A reprodução sexual é um processo mediante a qual dois organismos da mesma espécie unem seu material genético para dar lugar a um organismo fixo com combinação única de genes; para isso, cada organismo produz células que contém a metade do material genético característico da espécie. Essas células haploides (1n) são denominadas gametas; ao combinar-se um gameta masculino com um feminino produz-se uma célula diploide (2n) (zigoto ou ovo) a partir da qual se forma o embrião. A grande maioria das espécies com reprodução sexual são anisogâmicas, o que significa que produzem dois tipos de gametas diferentes: os gametas masculinos são microscópios, móveis e produzem-se em grande quantidade, enquanto que os femininos são grandes, imóveis e produzem-se em menor quantidade. O tipo de gameta que um indivíduo produz é o que define seu sexo; sobre os animais o macho é o indivíduo que produz grandes quantidades de espermatozoides e a fêmea produz uma menor quantidade de óvulos, enquanto que nas plantas as gônadas masculinas são as produtoras pólen e as femininas produzem oosferas. Os gametas são diferentes do resto das células do organismo, as quais se chamam células somáticas; essas últimas são diploides porque contém dois pares de cromossomos, um par herdado do pai do indivíduo e o outro da mãe. As células somáticas, ademais, se dividem por mitose, ao qual os cromossomos se duplicam antes de cada divisão celular e cada uma das células filhas recebe um complemento diploide idêntico dos cromossomos, logo todas as células somáticas de um indivíduo possuem o mesmo material genético, embora cada tipo celular expresse diferentes combinações de genes. Em contraponto, os gametas são células haploides porque possuem somente um par de cromossomos e a metade do material genético característico da espécie. Cada um dos cromossomos em um gameta é resultado da recombinação dos genes contidos nos cromossomos paterno e materno do indivíduo que originam o gameta, e cada um destes possuem uma combinação única de genes. Os gametas se formam a partir das células germinais, que são células que em sua origem são diploides e elas de “comprometem” a manter-se como uma linha celular especial que em determinado momento sofrerá o processo de meiose para dar origem aos gametas haploides, sejam óvulos ou espermatozoides segundo o sexo do animal. Como descrito no trabalho sobre a diferenciação sexual, as células germinativas primordiais originam-se no epiblasto do embrião, e migram desde o saco vitelino até colonizar as cristas gonodais, onde, por sua vez, proliferam-se e se organizam junto com as células somáticas da gônada primitiva para formar o testículo ou o ovário. As células germinais masculinas e femininas tem a mesma origem embrionária. As gônadas indiferenciadas em um embrião possuem três tipos celulares: as células que dão origem aos gametas (ovogonia ou espermatogonia), as precursoras de células que nutrem os gametas em desenvolvimento (células da granulosa no ovário; células de Sertoli no testículo) e as precursoras de células que secretam hormônios sexuais (células da teca no ovário; células de Leydig no testículo). As células germinais são as únicas estruturas do organismo que têm a capacidade de dividir-se por meiose sofrendo uma redução no número de seus cromossomos, sendo responsável pela transmissão da carga genética aos descendentes. Em contraste, as células somáticas somente se dividem por mitose. A formação dos gametas compreende fases sequenciais de mitose, meiose e pós-meiose. Esses processos são altamente organizados e necessitam de um preciso e bem coordenado programa de expressão genética. Uma das características importantes da gametogênese é a redução cromossômica, que através da meiose, reduz pela metade o número de cromossomos e produz células distintas entre si, devido a trocas de material genético entre os pares de cromossomos provenientes do pai e da mãe, o que ocorre no processo de “crossing over” durante a primeira fase da meiose. A gametogênese é o processo mediante o qual as células germinais de cada sexo se multiplicam, dividem e diferenciam até formar os gametas. No caso da formação dos gametas masculinos o processo recebe o nome específico de espermatogênese, e para os gametas femininos é denominado como ovogênese. Embora os dois processos alcancem o objetivo comum de produção das células haploides, por onde compartilham algumas características, existem diferenças marcadas entre eles devido a necessidade de produzir um número muito distinto de gametas, de tamanho diferente, e com características de motilidade também distintas. -/- •___ESPERMATOGÊNESE -/- A espermatogênese é o processo mediante o qual se produz os gametas masculinos denominados espermatozoides. Durante a vida fetal as células germinais e as células somáticas do testículo em formação organizam-se em túbulos seminíferos que se derivam dos cordões sexuais primários e conformam a maior parte da medula do testículo. Na etapa fetal cada tubo seminífero é delimitado por uma membrana basal, recoberta na parte interior pelas células precursoras das células de Sertoli (um tipo de células somáticas). No exterior do túbulo localizam-se as células precursoras das células de Leydig ou intersticiais (figura 1), que também são células somáticas. Entre a membrana basal e as células de Sertoli encontram-se algumas células germinais denominadas espermatogonias de reserva A0 (denominadas gonócitos) que serão o único tipo de células germinais presentes no testículo enquanto o animal não alcançar a puberdade. As células de Sertoli estabelecem na região basal uniões oclusoras entre si, formando parte da barreira hemato-testicular. As espermatogonias A0 localizam-se por dentro da membrana basal do túbulo seminífero, embora fora da barreira hemato-testicular. Figura 1: fase neonatal. Nota-se a grande infiltração de tecido intersticial em quase 50% da seção originando que os túbulos sejam pequenos e redondos em sua maioria. O citoplasma e núcleo das células pré-Leydig são notadas claramente por essa ser uma espécie suína onde o tecido intersticial está claramente diferenciado. Hematoxilina-eosina (X 220.5). Fonte: Embrapa. -/- O número de células de Sertoli no testículo depende da influência do hormônio folículo estimulante (FSH) presente durante a vida fetal e as primeiras etapas de vida pós-natal. A população de células de Sertoli ao chegar a puberdade se manterá fixa durante o resto da vida do animal; existe uma relação positiva entre o tamanho e a população de células de Sertoli e a capacidade de produção de espermatozoides do testículo. As células de Sertoli são as únicas células somáticas que estão no epitélio seminífero, e sua função é a nutrição, sustentação e controle endócrino das células germinais. As células de Sertoli participam ativamente no processo de liberação dos espermatozoides para a luz do túbulo. Nesse momento, as células de Sertoli realizam a fagocitose de parte do citoplasma do espermatozoide dos chamados corpos residuais. As células de Sertoli também fagocitam as células germinais que se degeneram no curso normal da espermatogênese. Essas células ainda sintetizam grande quantidade de proteínas, como por exemplo as proteínas ABP (androgen hinding protein), que transportam andrógenos para todo o aparelho reprodutivo, transferrinas, que transportam ferro para a respiração celular das células germinais e também às inibinas, que regulam a liberação de FSH pela hipófise, através de um sistema de retroalimentação (feedback) negativa (figura 2). Figura 2: epitélio seminífero, células de Sertoli (flecha) (400 X). Fonte: Embrapa. -/- Antes da puberdade dos túbulos seminíferos observam-se ao corte como estruturas de diâmetro pequeno, sem luz, e conformados unicamente pelas células de Sertoli e espermatogonias de reserva e rodeados por abundante tecido intersticial, ao que estão presentes as células precursoras das células de Leydig. Ainda antes da puberdade, a diferenciação celular manifesta-se primeiro pela presença de espermatócitos primários, os quais se degeneram em geral na fase de paquíteno, por falta de estimulação hormonal. A partir de que o animal chega a puberdade inicia-se o processo de espermatogênese, que se manterá durante toda a vida do animal, exceto em espécies de animais silvestres muito estacionais, ao qual pode se suspender durante a época não reprodutiva para voltar e ser retomada na época ou estação reprodutiva. Depois da puberdade, os túbulos seminíferos possuem um diâmetro muito maior; em seu interior observa-se um grande número de células germinais de todos os tipos, diferentes estádios de divisão, e em seu lúmen contém líquido e espermatozoides. Ainda sobre o alcancei da puberdade, as espermatogonias começam a dividir-se aceleradamente por mitose, enquanto que no espaço intersticial as células mesenquimais também começam a se diferenciar e a dar origem as células de Leydig (figura 3). A partir dessa etapa as células de Leydig (totalmente diferenciadas) são também evidentes no exterior do túbulo, junto com as células mioides ou peritubulares que o rodeiam o que ao contrair-se são responsáveis por controlar o avanço dos fluidos e as células presentes no lúmen do túbulo. As células mioides estão situadas ao redor do túbulo, e é creditado a elas a promoção da contração e da integridade estrutural do túbulo. Esse tipo celular apenas se diferencia na puberdade pela ação dos andrógenos (figura 4). As interações entre as células de Sertoli e as mioides parecem ter um papel importante na manutenção das funções do testículo. Durante o processo de espermatogênese, as espermatogonias de reserva dividem-se periodicamente e enquanto algumas células fixas permanecem como espermatogonias de reserva, outras proliferam e sofrem uma seção de divisões mitóticas durante as quais se vão diferenciando até formarem espermatócitos primários (espermatocitogênese ou fase de mitose), logo sofrem divisões especiais mediante as quais reduzem seu número de cromossomos (fase de meiose), e ao final trocam de forma para converter-se em espermatozoides (espermatocitogênese) (figura 5). Cada uma dessas etapas da espermato- gênese será descrito detalhadamente adiante, antes é necessário a explicação de algumas características das células de Sertoli e de Leydig que ajudarão a entender seu papel durante a espermatogênese. Figura 3: células de Leydig no espaço intersticial do testículo bovino adulto PAS (400 X). Fonte: Embrapa. -/- Figura 4: o estabelecimento da puberdade pela presença de espermatozoides no túbulo. Hematoxilina-eosina (400 X). Fonte: Embrapa. Figura 5: fases mitóticas das espermatogonias (A0 e B) para formação de um espermatócito primário e as duas fases de meiose que se sucedem antes da espermatogênese. Fonte: ZARCO, 2018. -/- Ao início da espermatocitogênese as uniões oclusoras entre as células de Sertoli se abrem por etapas (como as comportas de um submarino) para permitir a passagem das espermatogonias em direção ao centro do túbulo seminífero sem que se estabeleça uma continuidade entre o exterior e o interior da barreira hemato-testicular. Uma vez ultrapassada essa barreira, as sucessivas gerações de espermatogonias, espermatócitos, espermátides e espermatozoides irão se localizar em direção ao interior do túbulo seminífero, em estreita associação com as células de Sertoli. Em consequência, as células de Sertoli dividem o túbulo seminífero em dois compartimentos; o compartimento basal (debaixo das uniões oclusoras das células de Sertoli), ao qual residem as espermatogonias de reserva, e o compartimento adluminal (em direção ao centro do túbulo), cujos espaços entre as células de Sertoli desenvolvem o resto do processo de espermatogênese (meiose e espermatocitogênese). Esse feito é importante porque durante a vida fetal as únicas células germinais existentes eram as espermatogonias de reserva, pelo que os antígenos expressados por gerações mais avançadas (espermatogonias intermediárias, secundárias, espermátides e espermatozoides) não são reconhecidos como próprios do corpo pelo sistema imunológico. Logo, o anterior implica que deve existir uma barreira entre eles e o sangue para evitar um ataque imunológico. Em todas as etapas da espermatogênese, as células de Sertoli atuam como células de suporte para as células germinais, que sempre permanecem recoberta pela membrana das células de Sertoli. Também atuam como células nutricionais já que proporcionam o meio em que as células germinais se desenvolvem e maturam, assim como as substâncias que regulam e sincronizam as sucessivas divisões e transformações das células germinais. As células de Sertoli produzem hormônios, como estrógenos e inibina que atuam sobre a hipófise para regular a secreção das gonadotropinas que controlam a espermatogênese. As células de Leydig que residem no exterior do túbulo seminífero também são importantes para a espermatogênese: produzem a testosterona que estimula e mantém a espermatogênese, bem como serve como substrato sobre o qual atua como aromatizador das células de Sertoli para transformá-las em estrógenos. Como supracitado, para seu estudo podemos dividir a espermatogênese em três fase: espermatocitogênese, meiose e espermiogênese (figura 6). Agora, serão descritas cada uma dessas etapas. Em algumas espécies, incluindo no homem, os macrófagos representam o segundo tipo celular intersticial mais numeroso no testículo, depois das células de Leydig. Os macrófagos e vários subtipos de linfócitos são identificados nós testículos de ovinos e ratos. Eles estão intimamente associados com as células de Leydig e atuam juntamente na regulação da esteroidogênese. Figura 6: fluxograma da espermatogênese. -/- Espermatocitogênese -/- A espermatocitogênese, também chamada de etapa proliferativa ou de mitose, consiste numa série de divisões mitóticas sofridas pelas células descendentes de uma espermatogonia de reserva. Uma vez que a célula se divide, abandona o estado de reserva e começa um processo de diferenciação. As espermatogonias de reserva (denominadas espermatogonias A0 na rata ou As nos humanos) são células que existem desde a vida fetal e que permanecem mitoticamente inativas durante a infância. Uma vez que alcançam a puberdade começam a dividir-se em intervalos regulares, e as células filhas podem permanecer como espermatogonias de reserva ou abandonar a reserva e ingressar na dita espermatocitogênese. Uma vez abandonada a reserva, as células filhas que vão se formando em cada divisão permanecem unidas por pontes citoplasmáticas, constituindo um clone que se divide sincronicamente. As células que se formam depois de cada divisão continuam sendo espermatogonias, porém cada geração é ligeiramente diferente da anterior. Na rata, por exemplo, as espermatogonias tipo A0 ao dividir-se originam espermatogonias do tipo A1, que em sucessivas divisões formam espermatogonias dos tipos A2, A3 e A4, as quais, por sua vez, sofrem outra mitose para formar espermatogonias intermediárias e uma mais para formar espermatogonias do tipo B. Essas últimas se diferenciam (sem se dividir) em espermatócitos primários, processo em que termina a fase de espermatocitogênese, que literalmente significa processo de geração de espermatócitos. As espermatogonias tipo A0 são a fonte para a contínua produção de gametas. A metade delas se dividem e formam células iguais (as chamadas células tronco) e a outra metade forma as espermatogonias A1, que sofre novas divisões mitóticas e formam os tipos 2, 3 e 4. O tipo A4 sofre mitose para formar a intermediária (A In), que por mitose, forma a tipo B (figura 6). Esses tipos de espermatogonias podem ser identificadas em evoluções histológicas de acordo com sua organização topográfica na membrana basal dos túbulos seminíferos ou mediante seu conteúdo de heterocromatina. Outra maneira de diferenciação se baseia em marcadores moleculares específicos que distinguem as espermatogonias tronco (A0) das demais, com os fins de isolamento, desenvolvimento in vitro e transplante. As tipo B passam por mitose para formarem os espermatócitos primários; estes iniciam a primeira etapa da meiose formando os espermatócitos secundários; na segunda etapa da divisão meiótica, cada espermatócito secundário se divide e formam as chamadas espermátides. Quando o testículo alcança seu desenvolvimento total, a meiose completa-se e as espermátides originadas se convertem em espermatozoides. Um dos maiores sinais característicos desse fenômeno é o alargamento das espermátides e sua migração em direção ao lúmen do túbulo seminífero (figuras 4, 7 e 8). Figura 7: espermatogonias marcadas por imuno-histoquímica, anticorpo monoclonal TGFa (400 x). Figura 8: fases de divisões meióticas (M), espermatócitos em paquíteno (PA) e espermatócitos secundários (ES). -/- Figura 9: estádio posterior a liberação dos espermatozoides na luz do túbulo. Hematoxilina-eosina (400 x). Mediante as seis divisões mitóticas que ocorrem durante a espermatocitogênese se forma potencialmente um clone de 64 espermatócitos primários a partir de cada espermatogonia A que ingressa sobre o processo. Não obstante, algumas células sofrem apoptose em cada uma das etapas do processo, ao qual o número real formado é menor. Em outras espécies produzem-se um transcurso similar de divisões mitóticas sucessivas durante a espermatocitogênese, embora a nomenclatura utilizada possa ser distinta, por exemplo nos bovinos as duas últimas divisões mitóticas dão origem as espermatogonias de tipo B1 e B2. -/- Meiose -/- Uma vez que as espermatogonias B se diferenciam em espermatócitos primários, esses iniciam a etapa de meiose, com uma nova divisão; desta vez a divisão é do tipo meiótica. Ao completar-se a primeira divisão meiótica (meiose I) se obtém os espermató-citos secundários, que ao sofrer a segunda divisão meiótica (meiose II) dão origem as espermátides. Vale salientar que a meiose é o processo mediante o qual reduz-se a metade do número de cromossomos, pelo que as espermátides que se obtém são células haploides (1n). Os espermatócitos secundários que se formam depois da primeira divisão meiótica contém a metade do número normal de cromossomos, porém a mesma quantidade de DNA já que cada cromossomo é duplo. As espermátides formadas na conclusão da segunda divisão meiótica (figura 7), por sua vez, contém a metade dos cromossomos, e esse já não são duplos, já que se trata de células 1n. Também deve-se enfatizar que durante a meiose é relevante o entrecruzamento dos cromossomos homólogos, pelo que cada espermátide possui uma combinação única e diferente de genes paternos e maternos. Outro ponto que deve ser levado em consideração é que cada espermátide somente possui um cromossomo sexual; a metade das espermátides contém o cromossomo X herdado da mãe do macho que está levando a cabo a espermatogênese e a outra metade contém o cromossomo Y herdado de seu pai. Para cada espermatócito primário que entra no processo de meiose obtém-se cerca de quatro espermátides, pelo qual ao ser completada a meiose potencialmente se poderiam formar até 256 espermátides por cada espermatogonia que abandona a reserva e ingressa na espermatocitogênese. -/- Espermiogênese -/- Durante a espermiogênese, também chamada de fase de diferenciação, as esper-mátides sofrem, sem se dividir, uma metamorfose que as transforma em espermatozoides, os quais finalmente são liberados das células de Sertoli em direção ao lúmen do túbulo seminífero. A espermiogênese é um processo complicado e longo já que a espermátide deve sofrer complexas trocas nucleares, citoplasmáticas e morfológicas que resultam na forma-ção dos espermatozoides. Algumas dessas mudanças incluem a condensação do material nuclear para formação de um núcleo plano e denso, a eliminação do citoplasma para a constituição de uma célula pequena, a formação de uma estrutura especializada denomi-nada acrossomo ou tampa cefálica, e a formação do pescoço e da cauda (flagelo) do esper-matozoide, do que depende a sua motilidade. Durante a maior parte da espermiogênese, as espermátides se mantém com uma estreita associação com as células de Sertoli; logo, chega-se a observar, então, flagelos que se projetam em direção a luz do túbulo que pare-cem sair das células de Sertoli, sendo na realidade os flagelos dos espermatozoides que ainda não tinham sido liberados pelo lúmen. Ao liberar os espermatozoides em direção a luz do túbulo, as células de Sertoli realizam a fagocitose de parte do citoplasma dos espermatozoides (corpos residuais). Também fagocitam os restos de todas as células germinais que sofrem apoptose ou degeneram-se durante a espermatogênese. Credita-se que ao realizar essas funções as células de Sertoli podem fazer uma monitoração eficiente da espermatogênese, o que lhes permitiria emitir sinais para colaborar na regulação desse processo em nível gonodal e a nível sistêmico através da secreção de hormônios como a inibina e o estradiol. Além da inibina e activina, as células de Sertoli sintetizam outras proteínas, como a ABP (proteína ligadora de andrógenos) que serve como uma molécula de transporte de andrógenos dentro dos túbulos seminíferos, ductos deferentes e epidídimo, ou a transfer-rina, que transporta o ferro necessário para a respiração celular. -/- Resultados da espermatogênese -/- O resultado da espermatogênese não significa apenas uma simples multiplicação das células germinais (até 256 espermatozoides a partir de cada espermatogonia A1), senão que através dela são produzidos gametas haploides pequenos, móveis e com grande diversidade genética entre eles, ao mesmo tempo que se mantêm uma reversa de células mãe (espermatogonias A0) a partir das quais se poderiam originar novos ciclos de esper-matogênese durante o resto da vida do animal. -/- Controle hormonal da espermatogênese -/- Como mencionado, o FSH reproduz um importante papel para o estabelecimento das células de Sertoli durante a vida fetal e início da vida pós-natal. O começo da esper-matogênese também é estimulado pelo FSH, que atua sobre as células de Sertoli para estimular sua função e a ativação de sinais dessas células em direção as células germinais, incluindo-as a abandonar a reserva e ingressar na espermatogênese. O FSH, assim mesmo, estimula a mitose durante o resto da espermatogênese e aumenta a eficiência do processo, já que reduz a apoptose e a degeneração de espermatogonias intermediárias e do tipo B. O FSH também estimula as células de Sertoli para produzirem inibina e ABP. Uma vez iniciada a espermatogênese somente requerem níveis baixos de FSH para se mantê-la. As células de Sertoli também devem ser estimuladas pela testosterona para funcio-nar de maneira adequada; se requer também do LH hipofisário: hormônio que estimula as células de Leydig para produzir testosterona. Por sua vez, a secreção de LH e FSH é regulada pelo GnRH hipotalâmico: esse neurohormônio também faz parte do mecanismo de regulação da espermatogênese. A espermatogênese também é modulada em nível local mediante a produção de determinados fatores e interações entre as células. Dentro dos fatores locais podemos mencionar o fator de crescimento parecido com a insulina 1 (IGF-1), o fator de crescimen-to transformante beta (TGF- β), activina, ocitocina e diversas citocinas. Entre as intera-ções celulares existem tanto uniões de comunicação entre as células de Sertoli e as células germinais, como pontes citoplasmáticas entre todas as células germinais que formam o clone de células descendentes de uma espermatogonia A1. Uma vez que as células de Sertoli iniciam sua função na puberdade é possível manter experimentalmente a espermatogênese somente com testosterona, sem ser requeri-dos nenhum outro hormônio. A quantidade de espermatozoides produzidos, no entanto, é maior quando há presença do FSH. Abaixo do estímulo do FSH as células de Sertoli produzem estradiol e inibina, hormônios que geram uma retroalimentação sobre o eixo hipotálamo-hipofisário para a regulação da secreção de gonadotropinas. Em particular, a inibina reduz a secreção de FSH, pelo qual é factível que sirva como um sinal que evite uma excessiva estimulação as células de Sertoli. -/- Ciclo do epitélio seminífero -/- Em cada espécie as espermatogonias de reserva iniciam um novo processo de divi-sões celulares em intervalos fixos: a casa 14 dias no touro; 12 dias no garanhão e a cada 9 dias no cachaço (reprodutor suíno). A nova geração de células que começam a proliferar sobre a base do tubo deslocam-se em direção ao centro do túbulo a geração anterior, que por sua vez deslocam-se as gerações anteriores. Devido as mudanças que vão sofrendo cada geração celular se ajustam a tempos característicos de cada etapa, já que rodas as células em uma determinada seção do túbulo estão sincronizadas entre si pelas células de Sertoli; em cada espécie somente é possível encontrar um certo número de combinações celulares: 14 diferentes combinações no caso da rata, 8 no touro e 6 no ser humano. A sucessão de possíveis combinações até regressar a primeira combinação se conhece como o ciclo do epitélio seminífero. Na maioria das espécies os espermatozoides que são libera-dos em direção a luz do túbulo provém das células que entraram no processo de esperma-togênese quatro gerações antes que a geração que está ingressando nesse momento, pelo que a espermatogênese no touro dura ao redor de 60 dias e um pouco menos em outras espécies domésticas. Significa que os efeitos negativos das alterações na espermatogêne-se podem estar presentes até dois meses depois de que se produziram essas alterações. Como supracitado, geralmente se observa a mesma combinação celular em toda a área de uma determinada secção transversal do túbulo seminífero. No entanto, se fizermos uma série de secções, observa-se que ao longo do túbulo há uma sucessão ordenada de combinações (a primeira em uma determinada secção; a segunda combinação na seguinte secção, e assim sucessivamente em secções subsequentes até regressar a primeira combi-nação. Teremos, então, que ao início da divisão das espermatogonias A1 se produz de forma sincronizada em uma secção do túbulo, e vai-se transmitindo como uma onda peristáltica as secções adjacentes. Esse processo é denominado como onda do epitélio seminífero e graças à esse túbulo seminífero sempre tem secções em todas as etapas da espermatogênese, com o que se alcança uma produção constante de espermatozoides. -/- Alterações da espermatogênese -/- Nas espécies estacionais a espermatogênese, como já mencionado, pode reduzir-se ou inclusive suspender sua atividade fisiológica durante a época não reprodutiva dessas espécimes, porém esse processo fisiológico não pode ser considerado como uma altera-ção. No entanto, a espermatogênese só pode ser alterada pelas enfermidades ou por fatores externos. A principal causa de alterações na espermatogênese é o aumento da temperatura testicular. Por isso, os testículos são localizados na saco escrotal e são “caídos” para fora do corpo como pode-se observar nos bovinos, caprinos, ovinos, caninos e no próprio homem. A temperatura testicular deve estar cerca de 2 a 6 °C abaixo da temperatura corporal normal. As células germinais masculinas são sensíveis ao calor, pelo qual na maioria dos mamíferos os testículos se encontram fora da cavidade abdominal e existe um sofisticado sistema de termorregulação para mantê-los a uma temperatura menor que a corporal. Se a temperatura corporal for elevada ou se os testículos permanecerem na cavidade abdominal, ou ainda se os sistemas termorreguladores do testículo sejam afetados por fatores inflamatórios como edema ou falta de mobilidade testicular dentro do escroto, a temperatura do tecido testicular aumentará e a espermatogênese sofrerá alterações proporcionais ao excesso de temperatura e a duração da elevação. A espermatogênese também pode ser afetada pela exposição a hormônios ou a outras substâncias. É possível que a causa mais comum (sobretudo no homem) seja o uso de esteroides anabólicos, que elevam a concentração de andrógenos na circulação, provo-cando um feedback negativo sobre a secreção de gonadotropinas. Ao deixar de estimular-se o testículo pelas gonadotropinas, este deixará de produzir testosterona, e as concentra-ções de andrógeno exógeno nunca alcançará as altíssimas concentrações de testosterona que normalmente estão presentes a nível do tecido testicular por ser o local onde se produz o hormônio. Também se supõe que diversas substâncias com propriedades estrogênicas derivadas de processos industriais (indústria dos plásticos, hidrocarbonetos etc.) e presentes no ambiente (fatores xenobióticos) podem ser responsáveis pelas alterações na espermatogênese em diversas espécies, entre as quais se inclui o ser humano. -/- • OVOGÊNESE E FOLICULOGÊNESE -/- A ovogênese é o processo seguido pelas células germinais da fêmea para a forma-ção dos óvulos, que são células haploides. Durante a vida fetal as células germinais proliferam-se no ovário por mitose, formando um grande número de ovogonias, algumas das quais se diferenciam em ovócitos primários que iniciam sua primeira divisão meiótica para deter-se na prófase da divisão. Somente alguns desses ovócitos primários retornarão e concluirão a primeira divisão meiótica em algum momento da vida adulta do animal, dando origem a um ovócito secundário e a um corpo polar. O ovócito secundário inicia a sua segunda divisão meiótica, a qual volta a ficar suspensa até receber um estímulo apropriado, já que somente os ovócitos secundários que são ovulados e penetrados por um espermatozoide retornam e concluem a segunda divisão meiótica, dando origem a um óvulo (figura 10). O processo de ovogênese é realizado dentro dos folículos ovarianos, que também tem que sofrer um longo transcurso de desenvolvimento e diferenciação denominado foliculogênese pelo que a ovogênese como tal realiza-se dentro do marco desse último processo. Por essa razão, na seguinte seção descreverei tanto a ovogênese como a folicu-logênese, e a relação que existe entre ambos. Figura 10: representação da ovogênese. Na etapa de proliferação, as células germinais se diferen-ciam por mitose. A meiose I se caracteriza por uma prófase prolongada, ocorrendo a duplicação do DNA. Nas duas divisões, que ocorrem antes da ovulação e depois da fertilização, a quantidade de DNA é reduzida a 1n, com o fim de que a fusão dos pronúcles (singamia) pós-fertilização, seja gerado um zigoto com um número de cromossomos de 2n (diploide). -/- Geração de ovócitos primários e folículos primordiais Tanto a ovogênese como a foliculogênese iniciam-se durante a vida fetal, quando as células germinais primordiais provenientes do saco vitelino colonizam a gônada primitiva e, junto com as células somáticas z organizam-se para a formação dos cordões sexuais secundários, que se desenvolvem principalmente no córtex do ovário. Nesse período, as células germinais que colonizaram o ovário sofrem até 30 divisões mitóticas, proliferando-se até formar milhares ou milhões de ovogonias, que inicialmente formam “ninhos” constituídos cada um deles por um clone de várias ovogonias que descendem da mesma célula precursora e que se mantêm unidas por pontes citoplasmáticas, sincronizan-do suas divisões mitóticas. Nessa etapa alcança-se a máxima população de células germinais no ovário, que antes de nascer se reduzirá drasticamente por apoptose. No ovário do feto humano chegam a haver até sete milhões de células germinais que ao nascimento se reduzem a dois milhões. Os ovários fetais do bovino, de maneira análoga, chegam a ter até 2.100.000 células germinais, que ao nascimento reduzem para 130.000 aproximadamente. A redução no número de ovogonias produz-se ao mesmo tempo que essas células, que vêm dividindo-se por mitose e estão agrupadas em ninhos, iniciam sua primeira divisão meiótica para se transformarem em ovócitos primários: células germinais que se encontram em uma etapa de suspensão (diplóteno) da prófase da primeira divisão meiótica. Nesse período produz-se uma grande proporção de células germinais; as células somáticas dos cordões sexuais, por sua vez, emitem projeções citoplasmáticas que separam a isolam os ovócitos primários sobreviventes, ficando cada um deles rodeados por uma capa de células aplanadas da (pré) granulosa. Ao mesmo tempo em que se forma uma membrana basal entre as células da granulosa e o tecido intersticial do ovário. Ao ovócito primário rodeado de uma capa de células da (pré) granulosa aplanadas e delimita-das por uma membrana basal denomina-se de folículo primordial (figura 11). Nas vacas os folículos primordiais bem formados já estão presentes nos ovários a partir do dia 90 da gestação. A maioria dos folículos primordiais com os que nasce uma fêmea se manterão inativos durante um longo tempo; muitos deles durante toda a vida do animal. Nos folículos primordiais inativos tanto os ovócitos primários como as células da granulosa conservam sua forma original e mantém um metabolismo reduzido estritamente ao mínimo necessário para manter-se viáveis. Por essa razão, ao realizar um corte histológico de qualquer ovário as estruturas mais numerosas que se observam serão os folículos primordiais. No entanto, cada dia da vida de um animal, inclusive desde a vida fetal, um certo número de folículos primordiais reiniciam seu desenvolvimento, e a partir desse momento um folículo exclusivamente pode ter dois destinos: o primeiro, prosseguir seu desenvolvi-mento até chegar a ovular, e o segundo (que é muito mais frequente) encontrar em algum momento condições inadequadas que fazem fronteira com ele para parar seu desenvolvi-mento, levando-o a sofrer atresia e degenerar até desaparecer do ovário. Figura 11: sequência da foliculogênese apresentando as diferentes estruturas que podemos encontrar em cada fase. Fonte: ZARCO, 2018. Culminação da ovogênese A ovogênese somente se completará quando um ovócito primário reinicia a meio-se; completa sua primeira divisão meiótica para formar um ovócito secundário e um primeiro corpo polar e, quando, finalmente sofrer uma segunda divisão meiótica para formar um óvulo e um segundo corpo polar. Os óvulos são as células 1n que constituem os gametas femininos, pouco numerosos, grandes e imóveis. A grande maioria dos ovóci-tos primários, como veremos mais adiante, nunca retomam a meiose e, em consequência, não chegam a formar ovócitos secundários, e muitos dos ovócitos secundários tampouco sofrem uma segunda divisão meiótica, pelo que não chegam a formar os óvulos. Ao longo da vida de uma fêmea, na maioria das espécies, menos de 0,1% dos ovócitos primários (um a cada mil) chega a terminar a ovogênese, dando origem a um óvulo. O supracitado deve-se a que a ovogênese somente pode retomar-se e ser completa-da em ovócitos primários que se encontram dentro dos folículos primordiais que (uma vez ativados) vão alcançando diversas etapas de seu desenvolvimento em momentos precisos aos que encontram as condições ideais de oxigenação, nutrição, vascularização e exposição a fatores parácrinos e a exposição a concentrações de hormônios que se requerem para que o folículo continue em cada etapa de seu desenvolvimento com o processo de foliculogênese até chegar a ovular. Qualquer folículo que não esteja nessas condições ao longo do desenvolvimento sofrerá degeneração e atresia, pelo que o ovócito primário em seu interior nunca chegará ao ponto em que pode retomar a primeira divisão meiótica. No que resta da presente seção revisaremos o processo de foliculogênese em cujo marco se desenvolve a ovogênese; havemos que tomar de conta que essa última se limita ao que ocorre nas células germinais (ovogonia, ovócito primário, secundário e óvulo), pelo qual depende intimamente do desenvolvimento do folículo de que essas células formam parte. Em um princípio a ativação do folículo primordial e o desenvolvimento folicular são independentes das gonadotropinas: não se conhecem os mecanismos precisos median-te os quais um folículo primordial se ativa e reinicia seu desenvolvimento, nem como se decide quais folículos, dentre as dezenas de milhares de ou centenas de milhares presentes em um ovário se reativarão em um dia em particular. A reativação trata-se de uma liberação de influência de fatores inibidores, já que os folículos primordiais se reativam espontaneamente quando cultivados in vitro, isolados do resto do tecido ovariano. Uma vez que um folículo primordial se ativa, inicia-se um longo processo de desenvolvimento que somente depois de vários meses (ao redor de cinco meses no caso dos bovinos) o levará a um estádio em que seu desenvolvimento posterior requer a presença das gonado-tropinas; daí que se diz que as primeiras etapas do desenvolvimento são independentes das gonadotropinas. Durante a fase independente de gonadotropinas, um folículo primordial que tenha sido ativado e tenha começado a crescer; passará primeiro para a etapa de folículo primá-rio, caracterizada por conter um ovócito primário que está rodeado, por sua vez, por uma capa de células da granulosa, que não são planas, e sim cúbicas. Depois, se o folículo continuar crescendo se transformará em um folículo secundário, ao qual as células da granulosa começam a proliferar (aumentando em número) e se organizam em duas ou mais capas que rodeiam o ovócito primário. Entre o ovócito e as células da granulosa que o rodeiam se forma nesta uma zona pelúcida; ainda assim o ovócito mantém contato direto com essas células, mediante o estabelecimento de pontes citoplasmáticas que atravessam a zona pelúcida. Através dessas pontes citoplasmáticas as células da granulosa podem passar nutrientes e informação ao ovócito primário. O volume e o diâmetro do ovócito primário aumentam ao mesmo tempo que as células da granulosa proliferam-se, para incrementar as capas ao redor do ovócito. De maneira gradual o citoplasma do ovócito primário aumenta até 50 vezes seu volume e a proliferação das células continua. Esses folículos que possuem cada vez mais células e portanto mais capas de células da granulosa se denominam folículos secundários. Para evitar confusões, há a necessidade de nomen-clatura ao qual o folículo vá mudando de nome de primordial a primário e logo, de secun-dário, a terciário, por sua vez, o ovócito que encontra-se em seu interior, a todo momento, segue sendo um ovócito primário. Durante a etapa dependente de gonadotropinas, os folículos secundários começam a formar um espaço cheio de líquido, o antro folicular, desse modo se convertem em folí-culos terciários. Com a utilização de outra nomenclatura, a formação do antro marca a transição entre folículos pré-antrais (sem antro) e folículos antrais (com antro). Em algum momento dessa transição entre folículo secundário e terciário, também aparece a depen-dência de folículos em direção as gonadotropinas, pelo qual somente podem seguir crescendo na presença do hormônio luteinizante (LH) e do hormônio folículo estimulante (FSH). Nos bovinos e em outras espécies (para seu estudo), os folículos antrais são dividi-dos em pequenos, médios e grandes. Embora todos eles possuam um antro folicular, dependendo do seu grau de desenvolvimento requerem diferentes concentrações de gona-dotropinas para continuar o crescimento. Os folículos antrais mais pequenos somente re-querem concentrações baixas de LH e FSH, pelo qual podem continuar crescendo em qualquer momento do ciclo estral inclusive em animais que não estão ciclando (fêmeas em anestro pré-puberal, gestacional, lactacional, estacional). Nas etapas posteriores os folículos antrais requerem primeiro concentrações elevadas de FSH, e nas etapas finais somente podem continuar crescendo na presença de pulsos frequentes de LH, pelo qual somente os folículos que encontram-se sob concentrações apropriadas desses hormônios podem seguir crescendo. Por essa razão, nos animais que se encontram em anestro de qualquer tipo somente é possível encontrar folículos antrais pequenos ou médios, segundo a espécie, e nos animais que se encontram ciclando (estro) o maior tamanho folicular encontrado em um determinado dia do ciclo dependerá das concentrações de FSH e LH presentes nesse momento e nos dias anteriores. Um folículo que chega ao estado máximo de desenvolvimento, conhecido como folículo pré-ovulatório, ao final, somente chegará a ovular se for exposto a um pico pré-ovulatório de LH. Como supracitado, cada dia na vida de uma fêmea inicia seu desenvolvimento um certo número de folículos; a grande maioria sofrem atresia, mas depois da puberdade em cada dia do ciclo estral um ou vários folículos vão encontrando ao longo do seu desenvol-vimento concentrações hormonais que lhes permite chegar na etapa de folículo pré-ovula-tório. Somente nestes folículos, e como consequência de um pico pré-ovulatório de LH, se reinicia e completa-se a primeira divisão meiótica do ovócito primário, produzindo duas células distintas. Uma delas é o ovócito secundário, que retém praticamente todo o citoplasma. Contém, assim mesmo, em seu núcleo um par de cromossomos duplos, a outra é o primeiro corpo polar, que é exclusivamente um núcleo com uma quantidade mínima de citoplasma. Na maioria das espécies ovula-se um ovócito secundário que se encontra, então, suspendido na segunda divisão meiótica. Esta segunda divisão meiótica somente reinicia-rá e completarar-se uma vez que o espermatozoide começa a penetrar sob o ovócito secundário. Ao concluir-se a divisão se forma o segundo corpo polar e completa-se a ovogênese com o qual se obtém o óvulo, célula 1n que constitui o gameta feminino. No entanto, o óvulo existe pouco tempo como tal, já que em poucos minutos/horas (depen-dendo da espécie) se produzirá a fusão do núcleo do mesmo (pró-núcleo feminino) com o do espermatozoide (pró-núcleo masculino), com o qual se completa a fertilização e se forma um novo indivíduo (o ovo ou zigoto). -/- Ondas foliculares -/- Como mencionado supra, todos os dias um determinado número de folículos pri-mordiais se ativam e começam a crescer, os quais crescem em um ritmo característico em cada espécie. Isso provoca que em qualquer momento existam nos ovários folículos pri-mordiais (que começam a crescer em alguns dias ou semanas), assim como folículos secundários em diversas etapas do desenvolvimento, os quais iniciaram seu desenvolvi-mento em semanas ou inclusive meses (segundo o grau de desenvolvimento atual). Também em qualquer momento poderá haver folículos antrais nas etapas iniciais de seu desenvolvimento (com antros que já se podem detectar em cortes histológicos mas não são visíveis macroscopicamente). Todos esses folículos chegaram até seu estado de de-senvolvimento atual (primário, secundário ou antral pequeno), independente da etapa do ciclo estral em que sejam observados ou encontrados. Nos bovinos, os folículos que chegam ao início da etapa antral iniciaram seu desenvolvimento cinco meses antes, e todavia requerem ao redor de 42 dias para chegar ao estado pré-ovulatório. Para continuar seu desenvolvimento, os folículos antrais pequenos devem encon-trar concentrações altas de FSH, que os estimulam para prosseguir o crescimento. Cada vez que se produz uma elevação nas concentrações de FSH, esse hormônio estimula o desenvolvimento de um grupo de folículos antrais pequenos, que começaram a crescer muito tempo antes e que o dia da elevação de FSH tenha alcançado o grau de desenvolvi-mento preciso para responder com eficiência a este hormônio, o qual atuará através de seus receptores nas células da granulosa para estimular a produção de estradiol, a secreção de inibina, a produção de líquido folicular e a proliferação das células da granulosa. Um grupo de folículos antrais pequenos é assim recrutado pelo FSH para acelerar seu cresci-mento e aumentar sua produção de estradiol e inibina (figura 12). Mediante um seguimento ultrassonográfico dos ovários é possível identificar pou-cos dias depois um certo número de folículos, que por haver sido recrutados começam um período de crescimento acelerado. Durante alguns dias vários folículos crescem juntos, porém depois um deles é selecionado para continuar crescendo, enquanto que o restante do grupo deixam de fazê-lo e terminam sofrendo atresia. Através da ultrassom é possível identificar o folículo selecionado, agora chamado folículo domi-nante, já que sua trajetória de crescimento sofre um desvio com respeito a seguida pelo restante do grupo. Os folículos que não foram selecionados deixam de crescer e sofrem atresia já que deixam de possuir o suporte gonadotrópico de FSH, uma vez que as concentrações desse hormônio são suprimidos pela inibina e o estradiol produzidos pelo conjunto de folículos que conformam a onda folicular (figura 12), porém o folículo mais desenvolvido do grupo se converterá em dominante. A inibina atua diretamente a nível hipofisário para reduzir a secreção de FSH. Figura 12: onda folicular e relação dos níveis de FSH, estradiol e LH. Fonte: ZARCO, 2018. -/- Figura 13: Recrutamento, seleção e dominação folicular na espécie ovina e influência do FSH e LH nas fases. Fonte: SILVA, E. I. C. da, 2019. -/- A razão pela qual o folículo dominante é capaz de continuar seu desenvolvimento apesar da baixa nas concentrações de FSH é que o folículo é o único que alcançou o grau de progresso necessário para que apareçam os receptores para LH em suas células da granulosa. Esse processo permite ao folículo dominante ser estimulado pela LH, e que requeira baixas concentrações de FSH para manter seu desenvolvimento. A secreção de LH em forma de pulsos de baixa frequência (um pulso a cada quatro a seis horas), característica da fase lútea do ciclo estral; é suficiente para permitir que um folículo dominante continue crescendo por mais dias depois da sua seleção e que mais tarde mantenha-se viável durante alguns dias embora não aumentem de tamanho. Contu-do, se durante o período viável desse folículo não seja finalizada a fase lútea e não diminuam as concentrações de progesterona, o folículo terminará sofrendo atresia devido a exigência de um padrão de secreção acelerada de LH (aproximadamente um pulso por hora) durante o desenvolvimento pré-ovulatório, que somente pode ser produzido com a ausência da progesterona. Uma vez que um folículo dominante sofre atresia deixa de produzir inibina, pelo qual as concentrações de FSH podem elevar-se novamente para iniciar o recrutamento de outro grupo de folículos a partir da qual se origina uma nova onda folicular. Durante o ciclo estral de uma vaca podem gerar-se dois ou três ondas foliculares; somente em raros casos quatro. A etapa de dominância folicular da primeira onda na grande maioria dos casos não coincide com a regressão do corpo lúteo, pelo qual o primei-ro folículo dominante quase invariavelmente termina em atresia. Em algumas vacas o fo-lículo dominante da segunda onda ainda está viável quando se produz a regressão do corpo lúteo e acelera-se a secreção de LH, pelo qual esse segundo folículo dominante se converte em folículo pré-ovulatório e, ao final ovula. Em outros animais o segundo folícu-lo dominante também perde a sua viabilidade antes da regressão do corpo lúteo, por onde nesses animais se inicia uma terceira onda folicular, da qual surge o folículo que finalmen-te ovulará depois de produzir-se a regressão do corpo lúteo. Sem importar a onda em que se origine, uma vez que um folículo dominante é ex-posto a alta frequência de secreção de LH que se produz depois da regressão do corpo lúteo, aumenta ainda mais sua secreção de estradiol até que as altas concentrações desse hormônio comecem a exercer um feedback positivo para a secreção do LH. Isso provoca-rá a aceleração da frequência de secreção do LH até que os pulsos são tão frequentes que começam a ficar por cima e produzir-se o pico pré-ovulatório de LH, que é responsável pela realização da ovulação e a maturação final do ovócito. -/- •___DIFERENÇAS ENTRE ESPERMATOGÊNESE E OVOGÊNESE -/- Enquanto que na fêmea a ovogênese inicia-se durante a vida fetal, no macho a es-permatogênese começa na puberdade. Na fêmea, a partir de um ovócito primário se origi-na um óvulo; no macho, de um espermatócito primário se produzem, teoricamente, quatro espermatozoides. Outra característica interessante é que enquanto a fêmea já conta desde o nasci-mento com todos os ovócitos que necessitará na vida adulta, o macho necessitará chegar a puberdade para iniciar o desenvolvimento das células sexuais, já que ao nascer somente possui gonócitos precursores das células germinais, células de Sertoli e intersticiais. Na vida adulta de uma fêmea, o número de células germinais desaparece paulati-namente. Uma vez iniciada a espermatogênese no macho, a cada ciclo do epitélio seminí-fero as células germinais são renovadas mantendo a provisão para toda a vida reprodutiva. Na fêmea, a meiose sofre duas interrupções em seu transcurso, e no macho é ininterrupta. Figura 14: representação em diagramação comparativa do desenvolvimento da gametogênese. -/- Principais pontos abordados sobre as diferenças entre a gametogênese masculina e feminina: ❙ Na ovogênese a meiose contêm-se em duas ocasiões esperando acontecimentos externos para prosseguir. Já na espermatogênese não existe a suspensão da meiose. ❙ A espermatogênese é um processo contínuo, enquanto que a ovogênese pode completar exclusivamente um óvulo em cada ciclo estral; já que só pode ser completada por mais de um nas espécies que ovulam vários ovócitos no caso das porcas, cadelas, gatas etc. ❙ Na espermatogênese existem células de reserva que permitem a continuação du-rante toda a vida, enquanto que na ovogênese o número de ovócitos primários é limitado. A fêmea somente conta com os que nasceu, e eles não se dividem. ❙ Na espermatogênese obtém-se até 256 espermatozoides para cada espermatogo-nia que inicia o processo, enquanto que na ovogênese somente se obtém um óvulo a partir de cada ovócito primário. ❙ Durante a espermatogênese se produz uma metamorfose que transforma as es-permátides em espermatozoides. Na ovogênese não ocorre um processo análogo. ❙ Na espermatogênese, durante a meiose produzem-se quatro espermátides a partir de cada espermatócito primário. Na ovogênese se produz somente um óvulo a partir de cada ovócito primário; produz, ademais, dois corpos polares. ❙ Todos os óvulos que se produzem durante a ovogênese contém um cromossomo X, enquanto que a metade dos espermatozoides possuem um cromossomo Y e a outra metade um cromossomo X. ❙ Na espermatogênese produzem-se centenas ou dezenas de milhões de esperma-tozoides por dia, enquanto que na ovogênese se produz um ou alguns óvulos a cada ciclo estral. ❙ A espermatogênese produz gametas macroscópicos e com motilidade própria, enquanto que a ovogênese produz gametas grandes e imóveis. -/- REFERÊNCIAS BIBLIOGRÁFICAS -/- ABDEL-RAOUF, Mohammed et al. The postnatal development of the reproductive organs in bullswith special reference to puberty.(Including growth of the hypophysis and the adrenals). Acta endocrinologica, n. Suppl No. 49, 1960. ADONA, Paulo Roberto et al. Ovogênese e foliculogênese em mamíferos. Journal of Health Sciences, v. 15, n. 3, 2013. AERTS, J. M. J.; BOLS, P. E. 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Fisiologia Clínica do Ciclo Estral de Vacas Leiteiras: Desenvolvimento Folicular, Corpo Lúteo e Etapas do Estro. 2020. Acervo pessoal. DA SILVA, Emanuel Isaque Cordeiro. Fisiologia da Reprodução Animal: Ovulação, Controle e Sincronização do Cio. 2020. Acervo pessoal. DUKES, Henry Hugh; SWENSON, Melvin J.; REECE, William O. Dukes fisiologia dos animais domésticos. Editora Guanabara Koogan, 1996. EPIFANO, Olga; DEAN, Jurrien. Genetic control of early folliculogenesis in mice. Trends in Endocrinology & Metabolism, v. 13, n. 4, p. 169-173, 2002. ERICKSON, B. H. Development and senescence of the postnatal bovine ovary. Journal of animal science, v. 25, n. 3, p. 800-805, 1966. REFERÊNCIAS BIBLIOGRÁFICAS -/- FELDMAN, Edward C. et al. Canine and feline endocrinology-e-book. Elsevier health sciences, 2014. FUSCO, Giuseppe; MINELLI, Alessandro. The Biology of Reproduction. Cambridge University Press, 2019. GALINA-HIDALGO, Carlos Salvador. 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HYTTEL, Poul; SINOWATZ, Fred; VEJLSTED, Morten. Embriologia veterinária. São Paulo: Elsevier Brasil, 2012. JOHNSON, Martin H. Essential reproduction. Nova Jersey: John Wiley & Sons, 2018. JONES, Richard E.; LOPEZ, Kristin H. Human reproductive biology. Academic Press, 2013. KIERSZENBAUM, Abraham L.; TRES, Laura L. Primordial germ cell‐somatic cell partnership: A balancing cell signaling act. Molecular Reproduction and Development: Incorporating Gamete Research, v. 60, n. 3, p. 277-280, 2001. MATZUK, Martin M. et al. Intercellular communication in the mammalian ovary: oocytes carry the conversation. Science, v. 296, n. 5576, p. 2178-2180, 2002. MCLAREN, Anne. Germ and somatic cell lineages in the developing gonad. Molecular and cellular endocrinology, v. 163, n. 1-2, p. 3-9, 2000. MCKINNON, Angus O. et al. (Ed.). Equine reproduction. John Wiley & Sons, 2011. MERCHANT-LARIOS, Horacio; MORENO-MENDOZA, Norma. Onset of sex differentiation: dialog between genes and cells. 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Follicle growth in the immature mouse ovary. European Journal of Endocrinology, v. 62, n. 1, p. 117-132, 1969. PINEDA, Mauricio H. et al. McDonald's veterinary endocrinology and reproduction. Iowa state press, 2003. ROSER, J. F. Endocrine and paracrine control of sperm production in stallions. Animal Reproduction Science, v. 68, n. 3-4, p. 139-151, 2001. RUSSELL, Lonnie D. et al. Histological and histopathological evaluation of the testis. International journal of andrology, v. 16, n. 1, p. 83-83, 1993. RÜSSE, I.; SINOWATZ, F. Gametogenese. Lehrbuch der Embryologie der Haustiere, p. 42-92, 1991. SAITOU, Mitinori; BARTON, Sheila C.; SURANI, M. Azim. A molecular programme for the specification of germ cell fate in mice. Nature, v. 418, n. 6895, p. 293-300, 2002. SALISBURY, Glenn Wade et al. Physiology of reproduction and artificial insemination of cattle. WH Freeman and Company., 1978. SAWYER, Heywood R. et al. Formation of ovarian follicles during fetal development in sheep. 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Gametogénese. In. PORTA, L. R.; MEDRANO, J. H. H. Fisiología reproductiva de los animales domésticos. Cidade do México: FMVZ-UNAM, 2018. ZIRKIN, Barry R. et al. Endocrine and Paracrine Regulation of Mammalian Spermatogenesis. In: Hormones and Reproduction of Vertebrates. Academic Press, 2011. p. 45-57. -/- REALIZAÇÃO -/- . (shrink)

INTRODUÇÃO O anestro pós-parto é o período após o parto no qual a fêmea não apresenta ciclos estrais (atividade cíclica). Na vaca leiteira, o parto é seguido de um período de inatividade ovariana de duração variável, que é principalmente afetada pelo estado nutricional, produção leiteira, ganho ou perda de condição corporal antes e depois do parto, e por condições patológicas como hipoplasia dos ovários, cistos ovarianos, mumificação uterina, piometra entre outras, além, também, de condições ambientais como instalações que podem causar (...) estresse etc. 1. Controle neuroendócrino do anestro Entre os 12 e 15 dias pós-parto, as células gonadotróficas presentes na adenohipófise começam a produzir e secretar o FSH (hormônio folículo estimulante), o qual estimula a primeira onda folicular; entretanto, nenhum folículo dominante do(s) ovário(s) chega a completar seu desenvolvimento devido à carência de estímulo apropriado do LH (hormônio luteinizante) que também é produzido pelas mesmas células e que deve estar presente em altas concentrações. Nas 2 primeiras semanas pós-parto (primeiros 15 dias), as concentrações circulantes de LH são baixas, o que se deve à ausência de reservas do LH e à incapacidade temporal da hipófise para responder ao estímulo da GnRH (hormônio liberador de gonadotrofina). Entre os dias 15 e 20 pós-parto, a hipófise aumenta a sua capacidade para responder ao GnRH; entretanto, o estabelecimento do padrão de secreção do LH adequado para estimular a maturação e a ovulação do folículo dominante das primeiras ondas foliculares depende fundamentalmente do balanço energético. 2. Fatores que determinam o início da atividade ovariana pós-parto Na vaca leiteira, é frequente que algum dos primeiros folículos dominantes que se desenvolvem durante as primeiras duas ou três semanas complete a sua maturação e chegue à ovulação, isto é, madure e ovule. O fator limitante para o reinício da atividade ovariana é o balanço energético negativo (BEN). No gado leiteiro, o consumo de matéria seca (CMS), após o parto, deverá ser aumentado em até quatro vezes, para cobrir a demanda de nutrientes para a produção de leite e mantença; no entanto, a vaca é incapaz de consumir a exigência de matéria seca, por isso recorre a suas reservas de gordura e proteína, e cai em BEN. As vacas que consomem menos matéria seca, produzem menos leite, têm um BEN mais profundo e o período do parto à primeira ovulação é maior. O nível mais baixo do BEN é alcançado entre os 10 a 20 dias pós-parto e a vaca continua em BEN até os 70 a 80 dias pós-parto e, em alguns casos, até aos 100 dias pós-parto (vacas primíparas). Entretanto, mesmo com BEN, uma proporção elevada das vacas inicia sua atividade ovariana normal nas primeiras 8 semanas pós-parto. As vacas que alcançam um nível baixo de BEN rapidamente e saem deste ponto mínimo de forma rápida, iniciam sua atividade ovariana mais rápido também (20 a 30 dias pós-parto) do que aquelas que têm um BEN mais profundo e duradouro, essas muitas vezes permanecem em anestro aos 70 dias pós-parto (figura 1). As mudanças na condição corporal estão correlacionadas positivamente com as concentrações séricas de insulina, IGF-I e leptina; assim, a maior qualificação da condição corporal é maior mediante a concentração sérica desses hormônios, que atuam principalmente como sinais que chegam ao hipotálamo e modificam a frequência de secreção do GnRH e, consequentemente, de LH. A leptina é um hormônio que é produzido pelos adipócitos e tem sido proposto que sua concentração é o sinal mais importante de alterações de condição corporal. A transição do anestro para a ciclicidade coincide com um aumento das concentrações séricas de insulina, IGF-I e leptina. Além das mensagens dadas pelos hormônios supracitados, outras substâncias presentes no sangue fornecem informações sobre o estado metabólico do animal. Por exemplo, os ácidos graxos não esterificados e o β-hidroxibutirato são indicadores da mobilização e utilização da gordura corporal; assim, o aumento destas substâncias proporciona uma mensagem inibidora de reprodução. O intervalo médio entre o parto e a primeira ovulação, em vacas leiteiras em sistemas de produção não intensiva, é em torno de 30 dias; enquanto em vacas em sistemas de produção intensiva, é de 40 dias. Deve-se sinalar que é comum aos 70 dias pós-parto, que 20% das vacas ainda não ovulem. Verifica-se frequentemente que as vacas leiteiras desenvolvem folículos que crescem mais que os folículos ovulatórios, os quais convertem em cistos foliculares. Esta condição está associada com insensibilidade do hipotálamo ao retorno positivo do estradiol, o que não desencadeia o pico pré-ovulatório de LH (figura 2). 3. O corpo lúteo dos primeiros ciclos pós-parto Durante um ciclo estral o corpo lúteo tem uma meia-vida fisiológica de 12 a 14 dias; no entanto, na primeira ovulação pós-parto cerca de 25% das vacas desenvolvem corpos lúteos de vida curta (<10 dias), 30% de vida longa (21 a 50 dias) e 45% da vida normal (11 a 20 dias). A regressão prematura do corpo lúteo é uma condição frequente nos ruminantes no primeiro ciclo estral, da transição do anestro à ciclicidade. Ou seja, na vaca é frequente que no primeiro ciclo de pós-parto e da puberdade sejam ciclos curtos; isto deve-se à liberação antecipada da PGF2α. Por outro lado, a causa dos corpos lúteos de vida longa (corpos lúteos persistentes) não é clara, mas está relacionada com alterações na secreção da PGF2α devido a infecções uterinas (tabela 1). 4. Anestro patológico 4.1 Alterações do aparelho genital que causam anestro Na vaca foi calculado que as alterações do aparelho genital que afetam a atividade ovariana representam apenas 10% do total das causas de anestro. Entre as alterações do aparelho genital se encontram os cistos foliculares, cistos luteinizados, aplasia segmentar, piometra e mumificação fetal. . 4.2 Cistos foliculares Os cistos foliculares são a patologia ovariana mais comum nos bovinos leiteiros, causando perdas econômicas devido ao atraso do período do parto ao primeiro serviço, pelo custo dos tratamentos e pelo risco que têm as vacas de serem descartadas. A incidência aumentou à medida que a produção de leite se intensificou. Entre 5 e 30% das vacas desenvolvem cistos foliculares nos primeiros 60 dias do pós-parto; no entanto, cerca de 60% destas se recuperam espontaneamente. Sinais clínicos de vacas com cistos foliculares descritos na literatura são: ninfomania, ciclos curtos, masculinização e relaxamento dos ligamentos pélvicos. Atualmente uma alta proporção de vacas com cistos foliculares apresentam anestro. Por muitos anos um cisto folicular foi definido como um folículo com um diâmetro de 2,5 mm, presente em um ou em ambos os ovários durante um período mínimo de 10 dias, na ausência de um corpo lúteo. Os conhecimentos atuais modificaram o conceito clássico; assim, nem todos os cistos foliculares têm um diâmetro de 2,5 mm; além disso, alguns podem persistir mais de 10 dias. Outros são estruturas dinâmicas, que sofrem regressão e podem ser substituídas por novos cistos. Por esta razão, a definição mais precisa e técnica de um cisto folicular é: folículo de um diâmetro, de pelo menos 20 mm, que está presente em um ou ambos os ovários na ausência de tecido lúteo que interfere com o ciclo estral normal (figura 3). A patogênese dos cistos foliculares não é conhecida. Propõe-se que as vacas que desenvolvem esta patologia têm uma anomalia nos mecanismos de retroalimentação entre o hipotálamo e a hipófise, em conjunto com uma disfunção a nível folicular. É pesquisado e sugerido que em vacas com cistos foliculares, o pico pré-ovulatório de LH não ocorre ou é de menor amplitude, ou não tem uma relação síncrona com o amadurecimento do folículo, o que provoca a falha ovulatória. A alteração na secreção de LH pode obedecer a falta de sensibilidade do hipotálamo ao feedback positivo de estrogênio. Observa-se também que algumas anormalidades a nível folicular, tais como alterações na síntese de hormônios esteroides e uma menor sensibilidade à LH, podem contribuir com a patogenia. Uma vez que o cisto folicular foi estabelecido, observou-se um aumento na frequência de secreção pulsátil de LH, o que contribui com a persistência desta patologia. Existem fatores associados com a incidência de cistos foliculares que, de acordo com a patogenia proposta, podem influenciar ao nível do eixo hipotálamo-hipófise-ovário. Os cistos foliculares ocorrem principalmente durante a transição do anestro pós-parto para a ciclicidade. Neste período as vacas estão em BEN, e tem sido observado que as vacas que sofrem de um BEN mais profundo tendem a apresentar uma incidência maior de cistos foliculares. Por outra parte, há uma correlação positiva entre a produção de leite e a incidência de cistos, o que indica que as vacas que produzem mais leite têm maior risco de sofrer com esta patologia. Também existem outros fatores relacionados com a incidência de cistos foliculares tais como estresse, genéticos, infecções uterinas, estresse calórico e presença de fitoestrogênios na dieta (tabela 2). O tratamento indicado consiste na administração do GnRH, que provoca a luteinização do cisto. É recomendável a combinação com uma dose luteolítica de PGF2α, 7 a 10 dias depois da injeção de GnRH. Também é indicado o tratamento com hCG em vez de GnRH. Em alguns estudos foi possível integrar vacas com cistos foliculares em programas de sincronização da ovulação e inseminação em tempo fixo, com resultados aceitáveis de fertilidade (figura 4). 4.3 Cistos luteinizados Esse tipo de cisto pode se desenvolver a partir de um cisto folicular, também ocorre quando o saco libera um óvulo e, em seguida, se fecha novamente e enche-se de líquido. O tecido luteinizado produz progestágenos do qual os sinais incluem anestro persistente. São bastante mais raros e originam-se pelo contínuo aumento do espaço cavitário central normalmente existente nos corpos lúteos (figura 5). Normalmente, a parede do cisto contém células lúteogranulosas grandes e células tecaluteais pequenas. Se diferenciam com dificuldade de corpos lúteos recém-formados durante o período em que ainda possuem um antro. Os cistos luteínicos únicos, pelo menos os menores, necessitam de importância clínica. Se os cistos são funcionais, a sintomatologia mais destacada é o anestro ou pseudogestação. É preciso lembrar que esses cistos agem como uma fonte de secreção contínua de progesterona, diminuindo assim a resistência do útero a infecções, podendo desencadear-se uma endometrite (figura 6). Se eles forem diagnosticados é recomendável a administração de PGF2α. 4.4 Aplasia segmentar As vacas com esta condição podem gestar quando a ovulação ocorre do lado do corno uterino presente. Se a ovulação acontece do lado onde falta o corno uterino causa anestro. 4.5 Piometra A piometra, por definição, é a acumulação de material purulento no interior da luz uterina que ocorrem geralmente durante ou imediatamente após o período de domínio da progesterona. É uma condição que ocorre mais frequentemente nos primeiros 30 dias pós-parto; no entanto, algumas vacas em anestro que são revisadas no dia 50 pós-parto chegam a ter piometra. O tratamento indicado consiste na administração de PGF2α. 5. Anestro relacionado com o tempo de serviço O anestro em bovinos também é classificado como anestro pré-serviço e pós-serviço. 5.1 Anestro pré-serviço Inclui vacas e novilhas que não tenham mostrado estro, isto é, manifestado cio, no período em que devem ser servidas. Em alguns estudos observou-se que o anestro antes do serviço pode afetar até 50% das vacas elegíveis para apresentar estro. 5.1 Anestro pós-serviço Neste grupo incluem-se as vacas que não retornam ao estro, ou seja, que não retornaram ao cio, 21 dias depois da inseminação e não estão prenhes. Essas vacas são conhecidas como vacas fantasma (The Phantom Cow). A causa principal do atraso do retorno ao estro é a baixa eficiência na detecção de estros; mas também foram descritas outras causas como: inseminação em um estro anovulatório, inseminação em tempo fixo em vacas em anestro, suspensão da ciclicidade após o serviço, fases lúteas longas e morte embrionária. O diagnóstico precoce da gestação é de grande ajuda para conhecer oportunamente o número de fêmeas que não estão prenhes. A ecografia no trigésimo dia pós-inseminação adianta, pelo menos 10 dias, a ressincronização de vacas vazias. 6. Falso anestro ou anestro funcional Este causa as maiores perdas econômicas de caráter reprodutivo em rebanhos leiteiros, e refere-se às vacas que estão ciclando, mas não são observadas em estro pelos trabalhadores. É um dos maiores problemas dentro de um sistema de criação extensivo e, às vezes, semiintensivo, uma vez que é de suma importância a observação técnica e precisa do comportamento da vaca para observar o possível retorno ao cio ou não. 7. Manejo da vaca anéstrica A probabilidade de que a vaca seja inseminada oportunamente depende da eficiência da detecção de estros. É comum que a metade das vacas apresentem estro e não sejam observadas, por essa razão, muitas não são inseminadas uma vez que termina o período de espera voluntária. Por outro lado, há vacas que, por causas patológicas ou por seu estado metabólico, ainda estão em anestro depois que termina o período voluntário de espera. Para identificar as causas da ausência de estros, todas as vacas que não tenham sido inseminadas no dia 60 pós-parto, devem ser revisadas por via retal para aplicar o tratamento ou manejo pertinentes. Durante esta revisão é dada especial atenção às características do útero e nas estruturas ovarianas, já que daqui depende o manejo subsequente. A palpação começa no útero; neste é importante determinar se não há gestação. Posteriormente, avalia-se sua consistência que pode ser normal, edematosa ou turgente. Depois de avaliar o útero procede-se a palpação dos ovários, começando com o direito e, em seguida, esquerdo. Abaixo estão descritas as diferentes descobertas que podem ser encontradas na literatura e seu tratamento ou manejo. O registo dos resultados à palpação é feito através de chaves reprodutivas. 7.1 UN CLD2-3 FE10 Útero normal (UN) com um corpo lúteo 2 ou 3 no ovário direito (CLD2-3) e um folículo no ovário esquerdo de 10 mm de diâmetro (FE10). A consistência normal do útero (é normal quando não há edema ou turgescência) é encontrado em vacas não prenhes durante o diestro, ou em vacas que estão em anestro. O CL 2 ou 3 é uma estrutura bem desenvolvida que deforma o ovário e em alguns casos representa mais de 50% da massa ovariana. Classificá-los como CL 2 ou 3 é uma apreciação subjetiva do tamanho do corpo lúteo e não tem significado prático, pois em qualquer dos casos o manejo é o mesmo. O CL indica que a vaca está em qualquer dia do diestro e, obviamente, indica que está ciclando. Durante o diestro pode-se encontrar folículos de tamanho diferente em qualquer um dos dois ovários, já que este depende das ondas de desenvolvimento folicular. É importante frisar que as estruturas mencionadas podem estar em ovários diferentes ou no mesmo ovário. O achado mais importante nesta fase é a presença do corpo lúteo o que permite o tratamento com a PGF2α, o que resulta na apresentação do estro nas próximas 48 a 120 horas. A presença de um corpo lúteo é o estado fisiológico que é mais frequentemente encontrado neste grupo de vacas, primeiro porque paradoxalmente a maior proporção de vacas em “anestro” está ciclando (anestro funcional) e, em segundo lugar, porque o diestro ocupa 65% dos dias do ciclo estral. 7.2 UE CLD1 FE10-15 Útero edematoso (UE) com um corpo lúteo 1 no ovário direito (CLD1) e um folículo 10 ou 15 mm de diâmetro no ovário esquerdo. O útero edematoso pode ser encontrado no proestro e metaestro. A presença do CL1 e um folículo grande indica que se trata de uma vaca que muito provavelmente é encontrado em proestro. A diferença entre um CL1 e um CL 2 ou 3 é basicamente o seu tamanho; um CL1 é uma estrutura pequena com consistência dura. As vacas que têm estas características devem ser marcadas para que os trabalhadores prestem mais atenção, já que apresentarão o estro nas próximas 48 a 72 horas. Se a vaca não apresentar estro nesse período, deverá ser palpada na semana seguinte. 7.3 UT DE FE10-15 Útero túrgido ou tonificado, ovário direito estático e ovário esquerdo com um folículo de 10 ou 15 mm de diâmetro. Estes achados, além da presença de muco estral, correspondem a uma vaca em estro. Frequentemente, na palpação das vacas do grupo de anestro, há vacas em estro; estas vacas deverão ser programadas para inseminação. 7.4 UE DE EE Útero estático e ovários estáticos. Estas observações correspondem a uma vaca em metaestro; esta decisão tem uma alta margem de erro uma vez que também pode corresponder a um animal no proestro ou em anestro verdadeiro. Um achado que permite ser mais bem sucedido no diagnóstico é a presença de sangue no muco cervical; neste caso, a presença de sangue indica, com certeza, que a vaca está em metaestro; no entanto, nem todas as vacas apresentam este sangramento. As vacas com estes achados devem ser palpadas sete dias depois para confirmar ou corrigir um primeiro diagnóstico. Se a primeira palpação foi correta, na segunda se encontrará um CL2-3. 7.5 UE CHD FE10 Útero edematoso, com corpo hemorrágico no ovário direito (CHD) e com folículo de 10 mm de diâmetro no ovário esquerdo. Estas observações são de uma vaca em metaestro. O corpo hemorrágico é considerado como a fase de transição entre o folículo e o corpo lúteo; o CH é palpado como uma estrutura com uma saliência em forma de papila e é muito suave ao toque. Será necessário esperar 4 ou 5 dias para que se converta num corpo lúteo maduro e assim destruí-lo com PGF2α. Na rotina, estas vacas são palpadas na seguinte revisão (sete dias depois). 7.6 UN DE EE Útero normal e ovários estáticos. Este caracteriza as vacas que estão no anestro verdadeiro. As vacas caem no anestro principalmente por se encontrar em balanço negativo de energia; este problema é mais grave em vacas de primeiro parto. O único tratamento eficaz consiste em melhorar o seu estado metabólico. Tratamentos hormonais não funcionam caso não seja resolvido primeiro o seu estado nutricional. 7.7 UN CFD EE Útero normal, cisto folicular no ovário direito e ovário esquerdo estático. Embora as vacas com cistos foliculares sejam caracterizadas por apresentar estros recorrentes, também chegam a apresentar anestro. O tratamento consiste na administração do GnRH ou hCG. 7.8 UN CLD EE Útero normal, cisto luteinizado no ovário direito e ovário esquerdo estático. Este cisto também é causado por uma deficiência na secreção de LH, só que neste caso a deficiência foi parcial, o que provoca um certo grau de luteinização. O cisto luteinizado é uma estrutura com mais de 20 mm de diâmetro de paredes grossas. O tratamento indicado é a administração de PGF2α. Na prática, é difícil diferenciar um cisto folicular de um luteinizado, pelo que o tratamento recomendado é, primeiro, a administração de GnRH ou hCG e sete dias depois é injetado PGF2α (figura 7). 7.9 Piometra CLD2-3 FE A piometra é uma condição que ocorre principalmente nos primeiros 30 dias pós-parto; no entanto, na revisão das vacas anéstricas são encontradas vacas com esta patologia. O tratamento indicado consiste na administração de PGF2α. 8. Tratamentos hormonais para indução da atividade ovariana O estabelecimento precoce da ciclicidade pós-parto favorece a involução uterina e está positivamente correlacionado com a fertilidade; ou seja, quanto mais ciclos estrais a vaca tenha antes da primeira inseminação, a percentagem de concepção é maior. Um tratamento utilizado para induzir a primeira ovulação pós-parto em vacas anéstricas, consiste na administração de GnRH quando à palpação retal se encontra um folículo grande (> 10 mm). Com isso se pretende fazer o folículo ovular e depois injeta-se PGF2α para provocar a regressão lútea. Outras combinações consistem na administração de GnRH e na inserção de dispositivos de liberação de progesterona e, ao retirar o progestágeno, injeta-se PGF2α (figura 8). No entanto, os tratamentos mencionados são de uso comum, não funcionam em todos os casos. Um requisito para se obter êxito é que as vacas devem ter boa condição corporal ou que estejam ganhando condição corporal. Embora os tratamentos hormonais possam ajudar em alguns casos, não deve-se esquecer que as causas do anestro não são corrigidos apenas com a administração de hormônios, já que a vaca não cicla porque toda a informação do estado metabólico que recebe o cérebro, indica que não o deve fazer. 9. Gonadotropina coriônica equina (eCG) Na égua, por volta do dia 30 de gestação, células do córion migram para o endométrio e formam umas estruturas conhecidas como copas endometriais. Entre o dia 40 e o dia 130 de gestação nestas estruturas é produzido o eCG (anteriormente conhecida como gonadotropina sérica da égua prenha ou PMSG, pela sua abreviatura em inglês). Na égua esse hormônio tem atividade de LH pelo qual estimula a função do corpo lúteo e promove a formação de corpos lúteos acessórios. Nos ruminantes, o eCG se une aos receptores de LH e FSH do folículo, estimulando o desenvolvimento folicular, e no corpo lúteo estimula a secreção de progesterona. O eCG tem sido utilizado com a finalidade de superovulação e em vacas de corte sob anestro é incluído em programas com progestágenos para induzir ciclicidade. Nestes programas é injetado eCG no momento da remoção do progestágeno, o que favorece o desenvolvimento folicular e apresentação do estro. Em rebanhos leiteiros na Nova Zelândia tem sido incrementado a taxa de prenhez sem aumentar a proporção de partos gêmeos, através da injeção de eCG na remoção do dispositivo de libertação de progesterona em programas combinados com a injeção de benzoato de estradiol. RESUMO ➢ Transcorrem 15 dias pós-parto para que a hipófase responda ao hormônio GnRH. ➢ Entre os dias 12 e 15 pós-parto começa-se a secreção de FSH e iniciam as ondas foliculares. ➢ As vacas chegam ao ponto mais baixo de BEN entre os dias 15 e 20 pós-parto. ➢ As vacas iniciam a atividade ovariana normal, em média, após os 40 dias pós-parto. ➢ Aos 70 dias pós-parto é normal que 20% das vacas ainda estejam em anestro. ➢ Na primeira ovulação pós-parto 25% das vacas desenvolvem corpos lúteos de vida curta e 30% mostram corpos lúteos de vida longa (persistentes). ➢ As anormalidades do aparelho reprodutor são responsáveis por 10% das causas de anestro. ➢ 5 a 30% das vacas desenvolvem cistos foliculares nos primeiros 60 dias pós-parto e 60% destas se recuperam de forma espontânea. ➢ No dia 30 pós-inseminação é possível identificar vacas vazias mediante a ecografia e reservá-las para a ressincronização. ➢ A proteína B específica da gestação aparece no soro a partir do 15º dia de gestação. ➢ Para o diagnóstico precoce de gestação é recomendado medir a PSPB no dia 30 pósinseminação e em vacas com mais de 90 dias pós-parto. ➢ Entre os dias 50 e 60 pós-parto todas as vacas não servidas devem ser examinadas via retal para programar seu serviço. REFERÊNCIAS BIBLIOGRÁFICAS AMBROSE, Divakar J. Postpartum anestrus and its management in dairy cattle. Bovine Reproduction, p. 408-430, 2021. BALL, P. J. H.; PETERS, A. R. Reprodução em bovinos. São Paulo: SP, 2006. Características Gerais dos Bovinos/General Characteristics of Cattle Bovine. 2020. Fisiologia da Reprodução Animal: Ovulação, Controle e Sincronização do Cio. 2020. Nutrição Sobre a Reprodução e Fertilidade dos Bovinos. DERIVAUX, Jules; BECKERS, Jean-François; ECTORS, Francis. L'anoestrus du postpartum. Vlaams Diergeneeskundig Tijdschrift, v. 53, p. 215-229, 1984. ELLI, Massimiliano. Manual de reproducción en ganado vacuno. Zaragoza, ES: Servet Edit., 2005. FERNÁNDEZ DE CÓRDOBA DE LA BARRERA, Luis. Reproducción aplicada en el ganado bovino lechero. 1993. 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. GARVERICK, H. Allen. Ovarian follicular cysts in dairy cows. Journal of dairy science, v. 80, n. 5, p. 995-1004, 1997. HAFEZ, Elsayed Saad Eldin; HAFEZ, Bahaa. Reprodução animal. 2004. HIDALGO, Galina et al. Reproducción de animales domésticos. 2008. PETER, A. T.; VOS, P. L. A. M.; AMBROSE, D. J. Postpartum anestrus in dairy cattle. Theriogenology, v. 71, n. 9, p. 1333-1342, 2009. PETERS, A. R. et al. Reproducción del ganado vacuno. 1991. SPAIN, JAMES N.; LUCY, MATTHEW C.; HARDIN, DAVID K. Effects of nutrition on reproduction in dairy cattle. In: Current Therapy in Large Animal Theriogenology. WB Saunders, 2007. p. 442-450. UNGERFELD, Rodolfo. Reproducción de los animales domésticos. Edizioni LSWR, 2020. VILLA-GODOY, A. et al. Association between energy balance and luteal function in lactating dairy cows. Journal of Dairy Science, v. 71, n. 4, p. 1063-1072, 1988. WATHES, D. C. et al. Influence of negative energy balance on cyclicity and fertility in the high producing dairy cow. Theriogenology, v. 68, p. S232-S241, 2007. (shrink)

For Aristotle, a verb (ῥῆμα) is that which a) besides a proper meaning b) carry with it the notion of time; c) its parts do not significate separately and d) is a sign of something said of something else (OI ., 2, 16b6-8). This comprehensive definition distinguishes verbs from both nouns (since they do not carry the notion of time with themselves) and sentences or co-positings of words (since they have parts with independent meanings). Based on this definition, a verb (...) signifies three things: i) A main or proper meaning which is the meaning of the verb, e.g. running or sleeping; ii) time and iii) co-positing (or positing away). Although it is not asserted that one part is responsible for prossignification of time, its responsibility is given to co-positing-is. This might seem to be in contradiction with (c). However, it seems that (c) cannot include prossignifications of time and co-positing. What Aristotle means in (c) is just that parts of a verb cannot separately signify a meaning. Therefore, it is better to distinguish between signification, which is about meaning and prossignification, which is generally about anything but meaning. Hence, while the parts of the verb cannot have separate signification, they can have separate prossignification. Aristotle distinguishes between different kinds of verbs: 1) Definite verbs: those verbs that have definite nouns as their verb-name. 2) Indefinite verbs: those verbs that do not have definite nouns as their verb-name (OI., I, 3, 16b12-14; 10, 19b10-11). Aristotle’s examples of an indefinite verb are ‘is not-healthy’ and ‘is not-ill’. 3) Tenses of verbs: ‘was healthy’ and ‘will be healthy’ are tenses of the verb ‘is healthy’ because a verb indicates only the present time (OI., I, 3, 16b16-18). For Aristotle, almost all verbs either are composed of or can be analyzed to parts. An example of the former is ‘is healthy’ and of the latter is ‘runs,’ which can be analyzed to two parts: ‘is running.’ (OI., I, 10, 20a3-5; II, 12, 21b5-8). Thus, for him a verb either includes or can be analyzed to two parts: a part that includes the meaning and is, or is shown, by a name we call ‘verb-name’ (a), and a part that includes the co-positing of that meaning with the subject of the verb, which is or is shown by ‘is’, which we call ‘co-positing-is’ (d). The verb ‘runs’, for example, can be analyzed to a verb-name, ‘running’ and a co-positing-is (Note: Aristotle distinguishes between the verb ‘is’ in the sense of ‘exists’ and the co-positing ‘is’. Whereas in the former ‘is’ is a verb, having a signification of its own, which is ‘existing’, besides signifying time (OI., I, 10, 19b12-19), the ‘is’ in the latter is not a verb (Aristotle calls it ‘incidental’ is (OI., II, 11, 21a25-28) and is not sure whether to call it a verb or a noun (OI., I, 10, 19b22-24)) because it does not signify existing and, thus, need something else to be added. This something else is the meaning we are discussing. Therefore, it is only after the addition of the meaning to this ‘is’ that we have a verb. This is, I believe, the sense of the so much controversial text of OI, I, 3, 16b22-25: ‘For being and not being signify the existence of a thing not τὸ ὄν εἴπῃς αὐτὸ καθ᾿ ἑαυτὸ ψιλόν. For it does not indicate anything but prossignifies some co-positing (σύνθεσίν). Aristotle’s words at 16b24 about co-positing-is that it ‘prossignifies some co-positing’ (προσσημαίνει δὲ σύνθεσίν τινα) is an explicit evidence that she cannot be right.) and is not a concept apart from the things co-posited (συγκειμένων).’ This text, however, is so controversial. Originating from Ammonius and Boethius after him, who used the verb ‘copular’ for ‘est’ and Abelard, some commentators like Weidemann, Oehler, Francisco Ademollo take ‘is’ as having a copulative function. Allen Bāck notes that Aristotle does not separate off ‘is’ from other verbs as a mere copula but takes it ‘as the basic, paradigmatic form of the verb.’ He believes that in his interpretation of Aristotle (in: OI., 55, 19-24) Ammonius also took it as the paradigm for all verbs. Francisco Ademollo criticizes the standard view of taking ‘is’ as copula here and is inclined to take it as essential. David Ross thinks that although Aristotle is aware of the distinction between the existential and the copulative is, he has as yet no very clear idea of their relation. Although Jonathan Barnes agrees that the copula is surely present in de Interpretation, he thinks that at 16b22-25 Aristotle ought to be talking of existential εἶναι. Marie De Rijk Lambartus believes that there is no copula in Aristotle’s philosophical analysis of the elementary sentences and it is a mere corpus alienum in Aristotle’s semantics. Allan Bȃck believes that the copulative theory is not Aristotle’s and ‘may have come to dominate as a result of the neoPlatonizing interpretations of Aristotle’s works by Proclus, Ammonius and Boethius, which is the sign of the relation of running with a subject, e.g. Socrates in ‘Socrates runs’. Nonetheless, none of these parts significate separately. This is supposed to mean that a verb is a single signification. However, it is evident that each of the parts can signify if we use them in a different context as both of ‘running’ and ‘is’ in ‘runs’ or both of ‘is’ and ‘healthy’ in ‘is healthy’ can do. It might seem now that the first mentioned definition of verb might be contradictory. While Aristotle asserts that the parts of verb do not signify separately (d), he considers two other significations for a verb: the signification of time and relation. As it is obvious, both of these significations are done by one part of the verb, that is the co-positing-is. Therefore, in a verb e.g. ‘is healthy’ we have three significations: the significations of meaning, time and co-positing. Aristotle uses prossignification both for time (προσσημαῖνον χρόνον, OI., 2, 16b4) and co-positing (προσσημαίνει δἑ σύνθεσὶν τινα, OI, I, 3, 16b23-24). Since the copula part of a verb has no signification and does not form any conception, a verb is indeed its verb-name. Thus we see Aristotle saying that ‘verbs are in and by themselves names and signify something’ and like names, he adds, they arrest the hearer’s mind and fix his attention (OI., I, 3, 16b19-22). We can conclude then that verbs are names said of something else and carrying time. In other words, verbs are words that though have one single signification, as names have, they also have two other prossignifications: time and relation. (shrink)

Intra-molecular connectivity (that is, chemical structure) does not emerge from computations based on fundamental quantum-mechanical principles. In order to compute molecular electronic energies (of C 3 H 4 hydrocarbons, for instance) quantum chemists must insert intra-molecular connectivity “by hand.” Some take this as an indication that chemistry cannot be reduced to physics: others consider it as evidence that quantum chemistry needs new logical foundations. Such discussions are generally synchronic rather than diachronic —that is, they neglect ‘historical’ aspects. However, systems of (...) interest to chemists generally are metastable . In many cases chemical systems of a given elemental composition may exist in any one of several different metastable states depending on the history of the system. Molecular structure generally depends on contingent historical circumstances of synthesis and separation, rather than solely or mainly on relative energies of alternative stable states, those energies in turn determined by relationships among components. Chemical structure is usually ‘kinetically-determined’ rather than ‘thermodynamically-determined.’ For instance, cyclical hydrocarbon ring-systems (as in cyclopropene) are produced only in special circumstances. Adequate theoretical treatments must take account of the persistent effects of such contingent historical events whenever they are relevant—as they generally are in chemistry. (shrink)

Standpoint theorists have long been clear that marginalization does not make better understanding a given. They have been less clear, though, that social dominance does not make ignorance a given. Indeed, many standpoint theorists have implicitly committed themselves to what I call the strong epistemic disadvantage thesis. According to this thesis, there are strong, substantive limits on what the socially dominant can know about oppression that they do not personally experience. I argue that this thesis is not just implausible but (...) politically pernicious; it is an excuse for ignorance and silence that stifles our ability to address many injustices. Moreover, I argue that if we are to avoid lending support to the SEDT while working within a standpoint theory framework, we must hold that the socially dominant can achieve marginalized standpoints. So, we must hold that men can achieve feminist standpoints, that white women (and men) can achieve black feminist standpoints, and so on. (shrink)

What sort of logic do we get if we adopt a supervaluational semantics for vagueness? As it turns out, the answer depends crucially on how the standard notion of validity as truth preservation is recasted. There are several ways of doing that within a supervaluational framework, the main alternative being between “global” construals (e.g., an argument is valid iff it preserves truth-under-all-precisifications) and “local” construals (an argument is valid iff, under all precisifications, it preserves truth). The former alternative is by (...) far more popular, but I argue in favor of the latter, for (i) it does not suffer from a number of serious objections, and (ii) it makes it possible to restore global validity as a defined notion. (shrink)

It has been hypothesized that our believing that, or its seeming to us as though, the world is in some way dynamical partially explains (and perhaps rationalizes) future-bias. Recent work has, in turn, found a correlation between future-bias and near-bias, suggesting that there is a common explanation for both. Call the claim that what partially explains our being both future- and near-biased is our believing/it seeming to us as though the world is dynamical, the dynamical explanation. We empirically test two (...) versions of the dynamical explanation. The first is the moving ego explanation—according to which it is our belief that the ego moves, or our phenomenology as of the ego moving, that jointly (partially) explains future- and near-bias. The second is the moving time explanation—according to which it is our belief that time robustly passes, or our phenomenology as of robust passage, which jointly (partially) explain future- and near-bias. We found no evidence in favour of either explanation. (shrink)

I focus on three mereological principles: the Extensionality of Parthood (EP), the Uniqueness of Composition (UC), and the Extensionality of Composition (EC). These principles are not equivalent. Nonetheless, they are closely related (and often equated) as they all reflect the basic nominalistic dictum, No difference without a difference maker. And each one of them—individually or collectively—has been challenged on philosophical grounds. In the first part I argue that such challenges do not quite threaten EP insofar as they are either self-defeating (...) or unsupported. In the second part I argue that they hardly undermine the tenability of EC and UC as well. (shrink)

There is an undeniable tendency to dismiss women’s sexual assault allegations out of hand. However, this tendency is not monolithic—allegations that black men have raped white women are often met with deadly seriousness. I argue that contemporary rape culture is characterized by the interplay between rape myths that minimize rape, and myths that catastrophize rape. Together, these two sets of rape myths distort the epistemic resources that people use when assessing rape allegations. These distortions result in the unjust exoneration of (...) people we cannot conceive of as monstrous, while making it too easy to believe that some marginalized people could be rapists. I also argue that rape myths enable a novel kind of epistemic injustice. This injustice concerns how our assessments of trustworthiness and our assessments of plausibility interact. I argue that rape myths can result in runaway credibility deflations that can explain both why people fail to believe most women, and also why people may unjustly believe false allegations that white women have been raped by black men. (shrink)

We argue that there is a conflict among classical theism's commitments to divine simplicity, divine creative freedom, and omniscience. We start by defining key terms for the debate related to classical theism. Then we articulate a new argument, the Aloneness Argument, aiming to establish a conflict among these attributes. In broad outline, the argument proceeds as follows. Under classical theism, it's possible that God exists without anything apart from Him. Any knowledge God has in such a world would be wholly (...) intrinsic. But there are contingent truths in every world, including the world in which God exists alone. So, it's possible that God contingently has wholly intrinsic knowledge. But whatever is contingent and wholly intrinsic is an accident. So, God possibly has an accident. This is incompatible with classical theism. Finally, we consider and rebut several objections. (shrink)

Aristotle is essentially human; that is, for all possible worlds metaphysically consistent with our own, if Aristotle exists, then he is human. This is a claim about the essential property of an object. The claim that objects have essential properties has been hotly disputed, but for present purposes, we can bracket that issue. In this essay, we are interested, rather, in the question of whether properties themselves have essential properties (or features) for their existence. We call those who suppose they (...) do “property essentialists”; those who do not, “property anti-essentialists,” or “quidditists.” We offer two complementary arguments. Our total argument is under-girded by two assumptions: transworld identity theory and “received view” counterfactual semantics, a la David Lewis. We then argue that, if one presumes that these are true, then one risks running headlong into paradox if one also accepts property anti-essentialism. That's the first argument. By contrast, if one accepts these same assumptions in conjunction with property essentialism, then the paradox is avoided. This is the second argument. We take it that our arguments work to show that, between property essentialism and quidditism, the property essentialist is on better footing. Plausibly, properties themselves do have essential properties for their existence. (shrink)

Modal collapse arguments are all the rage in certain philosophical circles as of late. The arguments purport to show that classical theism entails the absurdly fatalistic conclusion that everything exists necessarily. My first aim in this paper is bold: to put an end to action-based modal collapse arguments against classical theism. To accomplish this, I first articulate the ‘Simple Modal Collapse Argument’ and then characterize and defend Tomaszewski’s criticism thereof. Second, I critically examine Mullins’ new modal collapse argument formulated in (...) response to the aforementioned criticism. I argue that Mullins’ new argument does not succeed. Third, I critically examine a powers-based modal collapse argument against classical theism that has received much less attention in the literature. Fourth, I show why God’s being purely actual, as well God’s being identical to each of God’s acts, simply cannot entail modal collapse given indeterministic causation. This, I take it, signals the death of modal collapse arguments. But not all hope is lost for proponents of modal collapse arguments—for the death is a fruitful one insofar as it paves the way for new inquiry into at least two new potential problems for classical theism. Showing this is my paper’s second aim. (shrink)

In her 2004 book Carol Gould addresses the fundamental issue of democratizing globalization, that is to say of finding ways to open transnational institutions and communities to democratic participation by those widely affected by their decisions. The book develops a framework for expanding participation in crossborder decisions, arguing for a broader understanding of human rights and introducing a new role for the ideas of care and solidarity at a distance. Reinterpreting the idea of universality to accommodate a multiplicity of cultural (...) perspectives, the author takes up a number of applied issues, including the persistence of racism, cultural rights, women's human rights, the democratic management of firms, the use of the Internet to enhance political participation, and the importance of empathy and genuine democracy in understanding terrorism and responding to it. Accessibly written with a minimum of technical jargon this is a major contribution to political philosophy. (shrink)

We can see mereology as a theory of parthood and topology as a theory of wholeness. How can these be combined to obtain a unified theory of parts and wholes? This paper examines various non-equivalent ways of pursuing this task, with specific reference to its relevance to spatio-temporal reasoning. In particular, three main strategies are compared: (i) mereology and topology as two independent (though mutually related) chapters; (ii) mereology as a general theory subsuming topology; (iii) topology as a general theory (...) subsuming mereology. Some more speculative strategies and directions for further research are also considered. (shrink)

Logical inferentialism maintains that the formal rules of inference fix the meanings of the logical terms. The categoricity problem points out to the fact that the standard formalizations of classical logic do not uniquely determine the intended meanings of its logical terms, i.e., these formalizations are not categorical. This means that there are different interpretations of the logical terms that are consistent with the relation of logical derivability in a logical calculus. In the case of the quantificational logic, the categoricity (...) problem is generated by the finite nature of the standard calculi and one direction in which it can be solved is to strengthen the deductive systems by adding infinitary rules (such as the ω-rule), i.e., to construct a full formalization. Another main direction is to provide a natural semantics for the standard rules of inference, i.e., a semantics for which these rules are categorical. My aim in this paper is to analyze some recent approaches for solving the categoricity problem and to argue that a logical inferentialist should accept the infinitary rules of inference for the first order quantifiers, since our use of the expressions “all” and “there is” leads us beyond the concrete and finite reasoning, and human beings do sometimes employ infinitary rules of inference in their reasoning. (shrink)

We draw out significant consequences of a relatively popular theory of metaphysical modality—branching actualism—for cosmological arguments for God’s existence. According to branching actualism, every possible world shares an initial history with the actual world and diverges only because causal powers (or dispositions, or some such) are differentially exercised. We argue that branching actualism undergirds successful responses to two recent cosmological arguments: the Grim Reaper Kalam argument and a modal argument from contingency. We also argue that branching actualism affords a response (...) to one popular defense of the classic contingency argument. What results are new difficulties for several cosmological arguments arising from the metaphysics of modality. (shrink)

I argue that Universalism (the thesis that mereological composition is unrestricted) entails Extensionalism (the thesis that sameness of composition is sufficient for identity) as long as the parthood relation is transitive and satisfies the Weak Supplementation principle (to the effect that whenever a thing has a proper part, it has another part disjoint from the first).

Learner’s attitude towards modular distance learning catches uncertainties as a world crisis occurs up to this point. As self-learning modules (SLMs) become a supplemental means of learning in new normal education, this study investigated efficiency towards the learners’ attitude and performance. Specifically, the study described the learners’ profile and their attitude and performance towards SLMs. It also ascertained the relationship between the learner’s profile with their attitude and performance, as well as the relationship between attitude and performance relevant to SLMs. (...) A descriptive-correlational research design was employed in the study with 88 non-randomly chosen participants from secondary schools in West Philippines. The learners’ GWA (general weighted average) was used to determine their performance and a survey questionnaire for their profile and attitude. Frequency counts, percentage, and Spearman’s rank correlation coefficient were used to analyze the data gathered with the aid of Jamovi software. Results revealed a strong correlation between attitude and performance, but there was no link between learner’s attitude and performance as to the learner’s profile. It was affirmed that attitude is a compelling factor that is related with performance. Hence, this research has broader ramifications that can direct the Bureau of Learning Delivery to orient the module writers to further contextualize the learning materials that can compound learner’s disposition and academic performance. (shrink)

Epistemic paternalism is the thesis that a paternalistic interference with an individual's inquiry is justified when it is likely to bring about an epistemic improvement in her. In this article I claim that in order to motivate epistemic paternalism we must first account for the value of epistemic improvements. I propose that the epistemic paternalist has two options: either epistemic improvements are valuable because they contribute to wellbeing, or they are epistemically valuable. I will argue that these options constitute the (...) foundations of a dilemma: either epistemic paternalism collapses into general paternalism, or a distinctive project of justified epistemic paternalism is implausible. (shrink)

Aristotle’s theory of human happiness in the Nicomachean Ethics explicitly depends on the claim that contemplation (theôria) is peculiar to human beings, whether it is our function or only part of it. But there is a notorious problem: Aristotle says that divine beings also contemplate. Various solutions have been proposed, but each has difficulties. Drawing on an analysis of what divine contemplation involves according to Aristotle, I identify an assumption common to all of these proposals and argue for rejecting it. (...) This allows a straightforward solution to the problem and there is evidence that Aristotle would have adopted it. (shrink)

One logic or many? I say—many. Or rather, I say there is one logic for each way of specifying the class of all possible circumstances, or models, i.e., all ways of interpreting a given language. But because there is no unique way of doing this, I say there is no unique logic except in a relative sense. Indeed, given any two competing logical theories T1 and T2 (in the same language) one could always consider their common core, T, and settle (...) on that theory. So, given any language L, one could settle on the minimal logic T0 corresponding to the common core shared by all competitors. That would be a way of resisting relativism, as long as one is willing to redraw the bounds of logic accordingly. However, such a minimal theory T0 may be empty if the syntax of L contains no special ingredients the interpretation of which is independent of the specification of the relevant L-models. And generally—I argue—this is indeed the case. (shrink)

Neutral monism is a position in metaphysics defended by Mach, James, and Russell in the early twentieth century. It holds that minds and physical objects are essentially two different orderings of the same underlying neutral elements of nature. This paper sets out some of the central concepts, theses and the historical background of ideas that inform this doctrine of elements. The discussion begins with the classic neutral monism of Mach, James, and Russell in the first part of the paper, then (...) considers recent neo-Russellian versions in the second half. The chances for a revival of neutral monism are probably slight; its key ideas and starting points lie far from those in contemporary philosophy of mind. A better route might be through the philosophy of science and a deeper understanding of causation. (shrink)

Intentions are commonly conceived of as discrete mental states that are the direct cause of actions. In the last several decades, neuroscientists have taken up the project of finding the neural implementation of intentions, and a number of areas have been posited as implementing these states. We argue, however, that the processes underlying action initiation and control are considerably more dynamic and context sensitive than the concept of intention can allow for. Therefore, adopting the notion of ‘intention’ in neuroscientific explanations (...) can easily lead to misinterpretation of the data, and can negatively influence investigation into the neural correlates of intentional action.We suggest reinterpreting the mechanisms underlying intentional action, and we will discuss the elements that such a reinterpretation needs to account for. (shrink)

Deception sometimes results in nonconsensual sex. A recent body of literature diagnoses such violations as invalidating consent: the agreement is not morally transformative, which is why the sexual contact is a rights violation. We pursue a different explanation for the wrongs in question: there is valid consent, but it is not consent to the sex act that happened. Semantic conventions play a key role in distinguishing deceptions that result in nonconsensual sex (like stealth condom removal) from those that don’t (like (...) white lies). Our framework is also applicable to more controversial cases, like those implicated in so-called “gender fraud” complaints. (shrink)

I argue that the conjunction of perdurantism (the view that objects are temporally extended) and universalism (the thesis that any old class of things has a mereological fusion) gives rise to undesired complications when combined with certain plausible assumptions concerning the semantics of tensed statements.

Are there any bona fide boundaries, i.e., boundaries that carve at the joints? Or is any boundary —hence any object—the result of a fiat articulation reflecting our cognitive biases and our so-cial practices and conventions? Does the choice between these two options amount to a choice between realism and wholesome relativism?

Integrated Information Theory (IIT) identifies consciousness with having a maximum amount of integrated information. But a thing’s having the maximum amount of anything cannot be intrinsic to it, for that depends on how that thing compares to certain other things. IIT’s consciousness, then, is not intrinsic. A mereological argument elaborates this consequence: IIT implies that one physical system can be conscious while a physical duplicate of it is not conscious. Thus, by a common and reasonable conception of intrinsicality, IIT’s consciousness (...) is not intrinsic. It is then argued that to avoid the implication that consciousness is not intrinsic, IIT must abandon its Exclusion Postulate, which prohibits overlapping conscious systems. Indeed, theories of consciousness that attribute consciousness to physical systems, should embrace the view that some conscious systems overlap. A discussion of the admittedly counterintuitive nature of this solution, along with some medical and neuroscientific realities that would seem to support it, is included. (shrink)

In previous work I have argued that talk about negative events should not be taken at face value: typically, what we are inclined to think of as a negative event (John’s failure to go jogging) is just an ordinary, positive event (his going to the movie instead); it is a positive event under a negative description. Here I consider more closely the difficulties that arise in those cases where no positive event seems available to do the job, as with putative (...) cases of causation by omission. In particular, I elaborate on Helen Beebee’s idea that not all causal explanations are reports of causation. When we mention John’s failure to turn off the gas as an explanans of why there was an explosion, we do not say what caused the explosion. We do not mention any of the relevant causes. We just remark that one sort of event that was supposed to occur, and whose occurrence would have prevented the explosion, did not in fact occur. (shrink)

That parthood is a transitive relation is among the most basic principles of classical mereology. Alas, it is also very controversial. In a recent paper, Ingvar Johansson has put forward a novel diagnosis of the problem, along with a corresponding solution. The diagnosis is on the right track, I argue, but the solution is misleading. And once the pieces are properly put together, we end up with a reinforcement of the standard defense of transitivity on behalf of classical mereology.

Aristotle’s typical procedure is to identify something's four causes. Intentional action has typically been treated as an exception: most think that Aristotle has the standard causalist account, according to which an intentional action is a bodily movement efficiently caused by an attitude of the appropriate sort. I show that action is not an exception to Aristotle’s typical procedure: he has the resources to specify four causes of action, and thus to articulate a powerful theory of action unlike any other on (...) offer. (shrink)

Traditionally, logic has been the dominant formal method within philosophy. Are logical methods still dominant today, or have the types of formal methods used in philosophy changed in recent times? To address this question, we coded a sample of philosophy papers from the late 2000s and from the late 2010s for the formal methods they used. The results indicate that the proportion of papers using logical methods remained more or less constant over that time period but the proportion of papers (...) using probabilistic methods was approximately three times higher in the late 2010s than it was in the late 2000s. Further analyses explored this change by looking more closely at specific methods, specific levels of technical engagement, and specific subdisciplines within philosophy. These analyses indicate that the increasing proportion of papers using probabilistic methods was pervasive, not confined to particular probabilistic methods, levels of sophistication, or subdisciplines. (shrink)

A lot of work in metaphysics relies on linguistic analysis and intuitions. Do we want to know what sort of things there are or could be? Then let’s see what sort of things there must be in order for what we truthfully say to be true. Do we want to see whether x is distinct from y? Then let’s see whether there is any statement that is true of x but not of y. And so on. In this paper I (...) argue that this way of proceeding is full of traps and is bound to be pretty useless unless we already have a good idea of what sort of things there are, and of how we are going to count them. (shrink)

Literature in epistemology tends to suppose that there are three main types of understanding – propositional, atomistic, and objectual. By showing that all apparent instances of propositional understanding can be more plausibly explained as featuring one of several other epistemic states, this paper argues that talk of propositional understanding is unhelpful and misleading. The upshot is that epistemologists can do without the notion of propositional understanding.

Peter Simons has argued that the expression ‘the universe’ is not a genuine singular term: it can name neither a single, completely encompassing individual, nor a collection of individuals. (It is, rather, a semantically plural term standing equally for every existing object.) I offer reasons for resisting Simons’s arguments on both scores.

We lay out the fatalist’s argument, making sure to clarify which dialectical moves are available to the libertarian. We then offer a more robust presentation of Ockhamism, responding to obvious objections and teasing out the implications of the view. At this point, we discuss presentism and eternalism in more detail. We then present our argument for the claim that the libertarian cannot take Ockham’s way out of the fatalism argument unless she rejects presentism. Finally, we consider and dispense with objections (...) to our argument. In the end, it ought to be clear that the libertarian must make a choice between Ockham’s way out and presentism. (shrink)

Some environmental ethicists and economists argue that attributing infinite value to the environment is a good way to represent an absolute obligation to protect it. Others argue against modelling the value of the environment in this way: the assignment of infinite value leads to immense technical and philosophical difficulties that undermine the environmentalist project. First, there is a problem of discrimination: saving a large region of habitat is better than saving a small region; yet if both outcomes have infinite value, (...) then decision theory prescribes indifference. Second, there is a problem of swamping probabilities: an act with a small but positive probability of saving an endangered species appears to be on par with an act that has a high probability of achieving this outcome, since both have infinite expected value. Our paper shows that a relative concept of infinite value can be meaningfully defined, and provides a good model for securing the priority of the natural environment while avoiding the failures noted by sceptics about infinite value. Our claim is not that the relative infinity utility model gets every detail correct, but rather that it provides a rigorous philosophical framework for thinking about decisions affecting the environment. (shrink)

In this paper I argue that composer John Cage’s so-called ‘silent piece’, 4’33”, is music. I first defend it against the charge that it does not involve the organization of sound, which has been taken to be a necessary feature of music. I then argue that 4’33” satisfies the only other condition that must be met for it to be music: it bears the right socio-historical connections to its predecessors within its tradition (Western art music). I argue further that one (...) cannot understand the organized sound condition and the socio-historical condition separately and that understanding their interaction has theoretical benefits—not least of which is providing a groundwork for a more culturally inclusive philosophy of music. Finally, I consider a number of outstanding questions concerning the content of the organized sound condition for Western art music in the wake of 4’33”. (shrink)

In this paper we argue that Socrates is a cognitivist about emotions, but then ask how the beliefs that constitute emotions can come into being, and why those beliefs seem more resistant to change through rational persuasion than other beliefs.

A critical review of the main themes arising out of recent literature on the semantics of ordinary event talk. The material is organized in four sections: (i) the nature of events, with emphasis on the opposition between events as particulars and events as universals; (ii) identity and indeterminacy, with emphasis on the unifier/multiplier controversy; (iii) events and logical form, with emphasis on Davidson’s treatment of the form of action sentences; (iv) linguistic applications, with emphasis on issues concerning aspectual phenomena, the (...) telicity/atelicity distinction, the treatment of statives, and temporal quantification. (shrink)

Recent research has begun treating the perennial philosophical question, “what makes a person the same over time?” as an empirical question. A long tradition in philosophy holds that psychological continuity and connectedness of memories are at the heart of personal identity. More recent experimental work, following Strohminger & Nichols (2014), has suggested that persistence of moral character, more than memories, is perceived as essential for personal identity. While there is a growing body of evidence supporting these findings, a critique by (...) Starmans & Bloom (2018) suggests that this research program conflates personal identity with mere similarity. To address this criticism, we explore how loss of someone’s morality or memories influence perceptions of identity change, and perceptions of moral duties towards the target of the change. We present participants with a classic ‘body switch’ thought experiment and after assessing perceptions of identity persistence, we present a moral dilemma, asking participants to imagine that one of the patients must die (Study 1) or be left alone in a care home for the rest of their life (Study 2). Our results highlight the importance of the continuity of moral character, suggesting lay intuitions are tracking (something like) personal identity, not just mere similarity. (shrink)

Despite their popularity, relatively scant attention has been paid to the upshot of Bayesian and predictive processing models of cognition for views of overall cognitive architecture. Many of these models are hierarchical ; they posit generative models at multiple distinct "levels," whose job is to predict the consequences of sensory input at lower levels. I articulate one possible position that could be implied by these models, namely, that there is a continuous hierarchy of perception, cognition, and action control comprising levels (...) of generative models. I argue that this view is not entailed by a general Bayesian/predictive processing outlook. Bayesian approaches are compatible with distinct formats of mental representation. Focusing on Bayesian approaches to motor control, I argue that the junctures between different types of mental representation are places where the transitivity of hierarchical prediction may be broken, and I consider the upshot of this conclusion for broader discussions of cognitive architecture. (shrink)