In recent years, a proliferation of books about empathy, cooperation and pro-social behaviours (Brooks, 2011a) has significantly influenced the discourse of the life-sciences and reversed consolidated views of nature as a place only for competition and aggression. In this article I describe the recent contribution of three disciplines – moral psychology (Jonathan Haidt), primatology (Frans de Waal) and the neuroscience of morality – to the present transformation of biology and evolution into direct sources of moral phenomena, a process here named (...) the ‘moralization of biology’. I conclude by addressing the ambivalent status of this constellation of authors, for whom today ‘morality comes naturally’: I explore both the attractiveness of their message, and the problematic epistemological assumptions of their research programmes in the light of new discoveries in developmental and molecular biology. (shrink)

Debates in the philosophy of science typically take place around issues such as realism and theory change. Recently, the debate has been reformulated to bring in the role of experiments in the context of theory change. As regards realism, Ian Hacking’s contribution has been to introduce ‘intervention’ as the basis of realism. He also proposed, following Imre Lakatos, to replace the issue of truth with progress and rationality. In this context we examine the case of the vitalism — reductionism debate (...) in biology inspired by the works of Indian physicist-turned-biologist Jagadish Chandra Bose (1858–1937), in the early twentieth century. Both camps had their characteristic hardcores. Vitalists led by John S. Burdon-Sanderson and Augustus D. Waller accepted religious metaphysics to support their research programme, which ultimately degenerated. Bose worked more with the ideals of science such as Occam’s razor, large-scale systematization of phenomena and novel prediction. I argue that his religious metaphysics, instead of acting as a protective shield, helped him to consolidate his position and allowed further problem shift resulting in a research programme that involved consciousness too. His research programme remains relevant even today. (shrink)

The generalized Darwinian research programme accepts physicalism, but holds that all life is purposive in character. It seeks to understand how and why all purposiveness has evolved in the universe – especially purposiveness associated with what we value most in human life, such as sentience, consciousness, person-to-person understanding, science, art, free¬dom, love. As evolution proceeds, the mechanisms of evolution themselves evolve to take into account the increasingly important role that purposive action can play - especially when quasi-Lamarckian evolution by cultural (...) means comes into existence. This programme of research brings together, into a coherent field of inquiry, aspects of such diverse fields of research as orthodox Darwinian theory (given its purposive interpretation), the study of animal behaviour, palaeontology, archaeology, history, anthropology, psycho-neurology, artificial intelligence, psychology, sociology, philosophy, linguistics, semantics, history and philosophy of science, and history and philosophy of inquiry more generally (the history and philosophy of ideas and culture). The great advantage of the generalized Darwinian research programme is that it provides a framework for understanding the deeds, achievements and experiences of people in a way that is compatible with the kind of knowledge and understanding achieved in the physical sciences, without being reducible to such knowledge and understanding. It promises to enable us to understand ourselves as a part of the biological domain without our humanity, our distinctive human value, being in any way denied: persons are not reduced to animals, and nor are animals misconceived to be persons. It holds out the hope that we can come to understand the human world as an integral part of the natural world without the meaning and value of the human world being thereby conceptually annihilated. The programme specifies in general terms what we must seek to do in order to develop a coherent understanding of nature and of ourselves which does justice to the character of both. (shrink)

“Teleosemantic” or “biosemantic” theories form a strong naturalistic programme in the philosophy of mind and language. They seek to explain the nature of mind and language by recourse to a natural history of “proper functions” as selected-for effects of language- and thought-producing mechanisms. However, they remain vague with respect to the nature of the proposed analogy between selected-for effects on the biological level and phenomena that are not strictly biological, such as reproducible linguistic and cultural forms. This essay (...) critically explores various interpretations of this analogy. It suggests that these interpretations can be explicated by contrasting adaptationist with pluralist readings of the evolutionary concept of adaptation. Among the possible interpretations of the relations between biological adaptations and their analogues in language and culture, the two most relevant are a linear, hierarchical, signalling-based model that takes its cues from the evolution of co-operation and joint intentionality and a mutualistic, pluralist model that takes its cues from mimesis and symbolism in the evolution of human communication. Arguing for the merits of the mutualistic model, the present analysis indicates a path towards an evolutionary pluralist version of biosemantics that will align with theories of cognition as being environmentally “scaffolded”. Language and other cultural forms are partly independent reproducible structures that acquire proper functions of their own while being integrated with organism-based cognitive traits in co-evolutionary fashion. (shrink)

Abstract: This study is designed to investigate the influence of menstruation on the academic achievement of senior secondary school Biology students in Irewole Local Government Area of Osun State, Nigeria. In the light of this, three specific objectives and research questions were highlighted not only to achieve but also to answer. The study employed quasi-experimental design. Besides, purposive sampling technique was used to take a sample of sixty-five (65) Grade 11 (Senior Secondary School II) students from three public high school (...) within the study area. The data generated from the instrument that is Biology Achievement Test (BAT) was analysed with the aid of a component of inferential statistics. The findings revealed a significant relationship between the academic achievement of students in Biology and menstruation. This is because, students before their menstrual period had mean score of 45 marks which was slightly higher than the one they had during (that is 42 marks) their menstrual period. A significant association between the academic achievement of students in Biology and menstrual pains was also observed. However, there was no significant relationship between the academic achievement in Biology among participants and duration of menstruation. This means that number of days used to complete menstruation period was insignificant to academic achievement of the participants in Biology. It is inferred from the study that there is relationship between menstruation and students’ academic performance in Biology among public secondary schools. Recommendations such as provision of conducive facilities in schools, constant awareness programme on the subject matter among others were made. (shrink)

One of the most influential arguments against the claim that computers can think is that while our intentionality is intrinsic, that of computers is derived: it is parasitic on the intentionality of the programmer who designed the computer-program. Daniel Dennett chose a surprising strategy for arguing against this asymmetry: instead of denying that the intentionality of computers is derived, he endeavours to argue that human intentionality is derived too. I intend to examine that biological plausibility of Dennett’s suggestion and (...) show that Dennett’s argument for the claim that human intentionality is derived because it was designed by natural selection is based on the misunderstanding of how natural selection works. (shrink)

This study is dedicated to the chapter Metaphysics My 10, which, in my opinion, represents an example of basic research as regards Aristotle’s ontological investigations. The aim of my analysis is to point out that the chapter constitutes a manifesto for a new ontology: this new ontology is Aristotle’s typological ontology. The main entities of the typological ontology are universals, on the one hand, and instantiations of universals, on the other hand. The ontological levels, to which these two kinds of (...) entities belong, are different from each other: therefore they must be kept rigorously separated from each other. The new ontology constitutes a clear opposition to the ontology of ideas. In my opinion Aristotle does not aim at eliminating universals from the range of the existing entities: At least the universals which correspond to biological kinds (such as „man“ or „animal“) and to all the properties connected to these biological kinds are real entities, even though they do not exist at the same level at which the level of the instantiations do. Aristotle’s aim is thus to assign universals and individuals to different realms of reality. The reality consists both of individuals and of universals: these kinds of entities occupy different realms of reality. (shrink)

The cumulative impact of enhancement technologies may alter the human species in the very long-term future. In this article, I will start showing how radical genetic enhancements may accelerate the conversion into a novel species. I will also clarify the concepts of ‘biological species’, ‘transhuman’ and ‘posthuman’. Then, I will summarize some ethical arguments for creating a transhuman or posthuman species with a substantially higher level of well-being than the human one. In particular, I will present what I shall (...) call the Principle of the Best Interests of Posthumanity, which states that the enhancement of the human and transhuman species must be directed towards the creation of a posthuman existence that is substantially more valuable than its predecessors. I suggest that human extinction may be considered, within that principle, as one of the best interests of posthumanity. Finally, I will develop three objections that make that principle unattractive and that show that pursuing a full-blown programme of posthuman evolution is ethically flawed. (shrink)

In a now classic paper published in 1991, Alberch introduced the concept of genotype–phenotype (G!P) mapping to provide a framework for a more sophisticated discussion of the integration between genetics and developmental biology that was then available. The advent of evo-devo first and of the genomic era later would seem to have superseded talk of transitions in phenotypic space and the like, central to Alberch’s approach. On the contrary, this paper shows that recent empirical and theoretical advances have only sharpened (...) the need for a different conceptual treat- ment of how phenotypes are produced. Old-fashioned metaphors like genetic blueprint and genetic programme are not only woefully inadequate but positively misleading about the nature of G!P, and are being replaced by an algorithmic approach emerging from the study of a variety of actual G!P maps. These include RNA folding, protein function and the study of evolvable soft- ware. Some generalities are emerging from these disparate fields of analysis, and I suggest that the concept of ‘developmental encoding’ (as opposed to the classical one of genetic encoding) provides a promising computational–theoretical underpinning to coherently integrate ideas on evolvability, modularity and robustness and foster a fruitful framing of the G!P mapping problem. (shrink)

The chapter argues from an ethology-inspired internalist perspective that ‘colour’ is not a homogeneous and autonomous attribute, but rather plays different roles in different conceptual forms underlying perception. It discusses empirical and theoretical evidence that indicates that core assumptions underlying orthodox conceptions are grossly inadequate. The assumptions pertain to the idea that colour is a kind of autonomous and unitary attribute. It is regarded as unitary or homogeneous by assuming that its core properties do not depend on the type of (...) ‘perceptual object’ to which it pertains and that‘colour per se’ constitutes a natural attribute in the functional architecture of the perceptual system. It is regarded as autonomous by assuming that it can be studied in isolation of other perceptual attributes. More generally, the chapter argues on the bases of corresponding empirical and theoretical evidence that perception cannot be understood as the ‘recovery’ of physical world structure from sensory structure by input-based computational processes. Rather, the sensory input serves as a kind of sign for biologically relevant aspects of the external world that activates biologically given conceptual forms with their internal parameters. Although the sensory input is a causally necessary requirement for the activation of conceptual forms, the perceptual computations triggered are under the control of an internal programme based on a set of conceptual forms. (shrink)

Historically, scientific and engineering expertise has been key in shaping research and innovation policies, with benefits presumed to accrue to society more broadly over time. But there is persistent and growing concern about whether and how ethical and societal values are integrated into R&I policies and governance, as we confront public disbelief in science and political suspicion toward evidence-based policy-making. Erosion of such a social contract with science limits the ability of democratic societies to deal with challenges presented by new, (...) disruptive technologies, such as synthetic biology, nanotechnology, genetic engineering, automation and robotics, and artificial intelligence. Many policy efforts have emerged in response to such concerns, one prominent example being Europe's Eighth Framework Programme, Horizon 2020, whose focus on “Responsible Research and Innovation” provides a case study for the translation of such normative perspectives into concrete policy action and implementation. Our analysis of this H2020 RRI approach suggests a lack of consistent integration of elements such as ethics, open access, open innovation, and public engagement. On the basis of our evaluation, we suggest possible pathways for strengthening efforts to deliver R&I policies that deepen mutually beneficial science and society relationships. (shrink)

The ecologist today finds scarce ground safe from controversy. Decisions must be made about what combination of data, goals, methods, and theories offers them the foundations and tools they need to construct and defend their research. When push comes to shove, ecologists often turn to philosophy to justify why it is their approach that is scientific. Karl Popper’s image of science as bold conjectures and heroic refutations is routinely enlisted to justify testing hypotheses over merely confirming them. One of the (...) most controversial theories in contemporary science is the Neutral Theory of Ecology. Its chief developer and proponent, Stephen Hubbell, presents the neutral theory as a bold conjecture that has so far escaped refutation. Critics of the neutral theory claim that it already stands refuted, despite what the dogmatic neutralists say. We see the controversy through a Popperian lens. But Popper’s is an impoverished philosophy of science that distorts contemporary ecology. -/- The controversy surrounding the neutral theory actually rests on a methodological fault. There is a strong but messy historical link between the concepts of being neutral and being null in biology, and Hubbell perpetuates this when he claims that the neutral theory is supplies the appropriate null for testing alternative theories. What method is being followed here? There are three contenders: -/- Null hypothesis testing tests for whether a there is a pattern to be explained. Null modeling tests for whether a process is causally relevant to a pattern. Baseline modeling apportions relative responsibility to multiple processes each relevant to a pattern. -/- Whether the neutral theory supplies an appropriate “null” depends upon whether null hypothesis, null modeling, or baseline model is intended. These methods prescribe distinct inference patterns. If they are null hypothesis testing or null modeling, the neutralists’s reasoning is invalid. If they are baseline modeling, the justification of a crucial assumption remains opaque. Either way, the neutral-null connection is being exploited rhetorically to privilege the neutral theory over its rivals. Clarifying the reasoning immunizes us against the rhetoric and foregrounds the underlying virtues of the neutralist approach to ecology. -/- The Popperian lens distorts theoretical development as dogmatism. Lakatos’s view of science as the development of research programmes clarifies the epistemology of the neutral theory. Focusing philosophical attention on the neutralist research programme illuminates (1) the synchronic uses of the neutral theory to make predictions and give descriptions and explanations; (2) its diachronic development in response to theoretical innovation and confrontation with data; (3) its complex relationships to alternative theories. For example, baseline modeling is now seen to be its primary explanatory heuristic. The justification for baseline modeling with the neutral theory, previously hidden from view, is seen in the logic of in the neutralist research programme. (shrink)

This paper discusses the role that appeals to theoretical simplicity have played in the debate between nativists and empiricists in cognitive science. Both sides have been keen to make use of such appeals in defence of their respective positions about the structure and ontogeny of the human mind. Focusing on the standard simplicity argument employed by empiricist-minded philosophers and cognitive scientists—what I call “the argument for minimal innateness”—I identify various problems with such arguments—in particular, the apparent arbitrariness of the relevant (...) notions of simplicity at work. I then argue that simplicity ought not be seen as a theoretical desideratum in its own right, but rather as a stand-in for other desirable features of theories. In this deflationary vein, I argue that the best way of interpreting the argument for minimal innateness is to view it as an indirect appeal to various potential biological constraints on the amount of innate structure that can wired into the human mind. I then consider how nativists may respond to this biologized version of the argument, and discuss the role that similar biological concerns have played in recent nativist theorizing in the Minimalist Programme in generative linguistics. (shrink)

In this book I show that science suffers from a damaging but rarely noticed methodological disease, which I call rationalistic neurosis. It is not just the natural sciences which suffer from this condition. The contagion has spread to the social sciences, to philosophy, to the humanities more generally, and to education. The whole academic enterprise, indeed, suffers from versions of the disease. It has extraordinarily damaging long-term consequences. For it has the effect of preventing us from developing traditions and institutions (...) of learning rationally devoted to helping us learn how to make progress towards a wiser, more civilized world. On our fragile earth, overcrowded, fraught with injustice, inequality and conflict, menaced by unprecedented and terrifying technological and industrial means for change and destruction, we urgently need to acquire a little more wisdom and civilization if we are to avoid repeating horrors of the kind suffered by so many during the 20th century (and already suffered by many in the first few years of the 21st century). We can ill afford to have in our hands instruments of learning botched and bungled from the standpoint of helping us learn how to live more wisely. It may seem scarcely credible that something as important as our institutions of learning can suffer from wholesale, structural defects. Why has this not been noticed before? Why have not armies of scientists and scholars appreciated the point and, long ago, put the matter right? The answer will emerge as my argument unfolds. I will show that one of the most damaging features of rationalistic neurosis is that it has built-in methodological and institutional mechanisms which effectively conceal that anything is wrong. But there is also an immediately obvious reason: specialization, and the resulting fragmentation of academia, has resulted in a situation in which hardly anyone takes responsibility for the overall ideals, the overall aims and methods, of academic inquiry. Academics, these days, are specialists, furiously trying to keep abreast of developments in their own specialist fields. They have no time, and no inducement, to lift their eyes from their particular disciplines, and look at the whole endeavour. Science has long been under attack, at least since the Romantic movement. Blake objected to “Single vision & Newton’s sleep” and declared that “Art is the Tree of Life... Science is the Tree of Death”. Keats lamented that science will “clip an Angel’s wings” and “unweave a rainbow”. Whereas the Enlightenment had valued science and reason as tools for the liberation of humanity, Romanticism found science and reason oppressive and destructive, and instead valued art, imagination, inspiration, individual genius, emotional and motivational honesty rather than careful attention to objective fact. Much subsequent opposition to science stems from, or echoes, the Romantic opposition of Blake, Wordsworth, Keats and many others. There is the movement Isaiah Berlin has described as the “Counter-Enlightenment” (Berlin, 1979, ch. 1). There is existentialism, with its denunciation of the tyranny of reason, its passionate affirmation of the value and centrality of irrationality in human life, from Dostoevsky, Kierkegaard and Nietzsche to Heidegger and Sartre (see, for example, Barrett, 1962). There is the attack on Enlightenment ideals concerning science and reason undertaken by the Frankfurt school, by postmodernists and others, from Horkheimer and Adorno to Lyotard, Foucault, Habermas, Derrida, MacIntyre and Rorty (see Gascardi, 1999). The soul-destroying consequences of valuing science and reason too highly is a persistent theme in literature: it is to be found in the works of writers such as D.H. Lawrence, Doris Lessing, Max Frisch, Y. Zamyatin. There is persistent opposition to modern science and technology, and to scientific rationality, often associated with the Romantic wing of the green movement, and given expression in such popular books as Marcuse’s One Dimensional Man, Roszak’s Where the Wasteland Ends, Berman’s The Reenchantment of the World and Appleyard’s Understanding the Present. There is the feminist critique of science and conceptions of science: see, for example, Fox Keller (1984) and Harding (1986). And there are the corrosive implications of the so-called “strong programme” in the sociology of knowledge, and of the work of social constructivist historians of science, which depict scientific knowledge as a belief system alongside many other such conflicting systems, having no more right to claim to constitute knowledge of the truth than these rivals, the scientific view of the world being no more than an elaborate myth, a social construct (see Barnes and Bloor, 1981; Bloor, 1991; Barnes, Bloor and Henry, 1996; Shapin and Schaffer, 1985; Shapin, 1994; Pickering, 1984; Latour, 1987). This latter literature has provoked a counter-attack by scientists, historians and philosophers of science seeking to defend science and traditional conceptions of scientific rationality: see Gross and Levitt (1994), Gross, Levitt and Lewis (1996), and Koertge (1998). This debate between critics and defenders of science came abruptly to public attention with the publication of Alan Sokal’s brilliant hoax article ‘Transgressing the boundaries’ in a special issue of the cultural studies journal Social Text in 1996 entitled Science Wars: see Sokal and Bricmont (1998). What does this book have to contribute, given the above avalanche of criticism of science? The criticisms of science developed in this book are diametrically opposed to the above, in so far as the above criticisms oppose scientific rationality, seek to diminish or restrict its influence, or hold that it is unattainable. My central point is that we suffer, not from too much scientific rationality, but from not enough. What is generally taken to constitute scientific rationality is actually nothing of the kind. It is rationalistic neurosis, a characteristic, influential and damaging kind of irrationality masquerading as rationality. Science is damaged by being trapped within a widely upheld but severely defective philosophy of science; free science from this defective philosophy, provide it with a more intellectually rigorous philosophy, and it will flourish in both intellectual and humanitarian terms. And more generally, as we shall see, academic inquiry as a whole is damaged by being trapped within an intellectually defective philosophy of inquiry; free it from this defective philosophy, from its rationalistic neurosis, and it will flourish in intellectual and human terms. It is not reason that is damaging, but defective pretensions to reason — rationalistic neurosis —masquerading as reason. Thus what I seek to do here is the exact opposite of what all those who oppose science and scientific rationality do. I shall argue that Reason, the authentic article, arrived at by generalizing the progress-achieving methods of science, can have profoundly liberating and enriching consequences for all worthwhile, problematic aspects of life, and thus deserves to enter into every aspect of life. The bare bones of the argument of this book can be stated quite simply like this. Science cannot proceed without making the substantial metaphysical assumption that the universe is physically comprehensible (to some extent at least). But this conflicts with the orthodox view that in science everything is assessed impartially with respect to evidence, nothing being permanently assumed independently of evidence. So the metaphysical assumption of comprehensibility is repressed. Science pretends that no such assumption is made. But this damages science. For the assumption is substantial, influential and highly problematic. It needs to be made explicit within science so that it can be critically scrutinized, so that alternatives can be developed and considered. Pretending the assumption is not being made undermines the intellectual rigour of science, its intellectual value and success. And it does not stop there. For science also makes value assumptions. Quite properly, science is concerned to discover that which is of value. New factual knowledge devoid of all value (whether intellectual or practical) does not contribute to science. But the orthodox view of science holds that values have no place within science. As in the case of metaphysics, here too, science pretends that values have no role to play within the intellectual domain of science. And this damages science. For values are, if anything, even more influential and problematic than metaphysics (in influencing the direction of research). Here too, values need to be made explicit within science so that they can be scrutinized, so that alternatives can be developed and assessed. Pretending that values play no role within the intellectual domain damages science; both the intellectual and the practical aspects of science are adversely affected. Science fails to pursue those avenues of research that lie in the best interests of humanity. And it goes further. Science is pursued in a social, cultural, economic and political context. It is a part of various social, economic and political projects which seek to achieve diverse human objectives. But the idea that science is an integral part of humanitarian or political enterprises with political ends clashes, once again, with the official view of science that the aim of science is to improve factual knowledge. The political objectives of science are repressed. And, once again, this damages science. For, of course, the political objectives of science, like all our political objectives, are profoundly problematic. These need to be made explicit so that they can be scrutinized, so that alternatives can be developed and considered. The pretence that science does not have this political dimension once again undermines the intellectual rigour of science, and its human value. It lays science open to becoming a part of economic, corporate and political enterprises that are not in the best interests of humanity. The upshot of the line of argument just indicated is that we need to bring about a revolution in the aims and methods of science, and of academic inquiry more generally. Natural science needs to change; its relationship with the rest of academic inquiry, with social inquiry and the humanities, needs to change; and most importantly and dramatically, academic inquiry as a whole needs to change. The basic task of the academic enterprise needs to become to help humanity learn how to tackle its problems of living in more cooperatively rational ways than at present. We need to put the intellectual tasks of articulating our problems of living, and proposing and critically assessing possible solutions, possible and actual actions, at the heart of academic inquiry. The basic task needs to be to help humanity learn a bit more wisdom – wisdom being the capacity to realize what is of value in life, for oneself and others, wisdom including knowledge and technological know-how, but much else besides. Natural science, despite its flaws, has massively increased our knowledge and technological know-how. This in turn has led to a massive and sometimes terrifying increase in our power to act. Often this unprecedented power to act is used for human good, as in medicine or agriculture. But it is also used to cause harm, whether unintentionally (initially at least) as when industrialization and modern agriculture lead to global warming, destruction of natural habitats and rapid extinction of species, or intentionally, as when the technology of war is used by governments and terrorists to maim and kill. Before the advent of modern science, when we lacked the means to do too much damage to ourselves and the planet, lack of wisdom did not matter too much. Now, with our unprecedented powers to act, bequeathed to us by science, lack of wisdom has become a menace. This is the crisis behind all the other current global crises: science without wisdom. In these circumstances, to continue to pursue knowledge and technological know-how dissociated from a more fundamental quest for wisdom can only deepen the crisis. As a matter of urgency, we need to free science and academia of their neuroses; we need to bring about a revolution in the academic enterprise so that the basic aim becomes to promote wisdom by intellectual and educational means. At present science and the humanities betray both reason and humanity. The argument of this book, which I have just summarized, begins with a discussion of the philosophical and methodological problems that beset theoretical physics. At once, many of those who are concerned about moral, political and environmental issues that plague our world today, will feel impatient. What have the methodological problems of theoretical physics to do with third world poverty, war and the threat of war, pollution, extinction of species, the menace of conventional, chemical, biological and nuclear armaments? I can only plead with such a reader: patience. The root cause of the sickness of our times does indeed lie, I claim, with a methodological sickness of natural science or, even more specifically, of physical science. What makes the modern world so utterly different from all previous ages is our possession of modern science and technology. This is the instrument that has changed the conditions of human life almost beyond all recognition, and put unprecedented, and sometimes terrifying, powers into our hands. One should not dismiss out of hand the suggestion that a part of our problem may lie with this instrument, this engine, of rapid change. We see, here, too, the way in which intellectual specialization, referred to above, has effectively concealed from view the nature and extent of the problem that confronts us. The argument that I shall develop in what follows begins with physics, with problems concerning the aims and methods of physics. But it then leaps to social science, to philosophy, to the humanities, to education, to psychotherapy, and to politics: in the end there is scarcely an aspect of modern life that is not touched and, potentially, affected by the argument. How many academic experts are prepared to follow, to take seriously, an argument that ranges so recklessly across such a wide range of academic disciplines? How many non-academic non-experts? But just this is what the argument and message of this book requires. I hope that the reader will endure with patience the somewhat esoteric discussion concerning the aims and methods of physics pursued in chapter one. This may not seem to have anything much to do with the urgent problems of our times, but it does. This is in part how the intellectual disaster I seek to expose in this book conceals itself from view: it buries itself in the obscure, recondite field of the philosophy of physics. Those who devote their lives to the philosophy of physics are, by and large, too myopic to see how their specialized field of study has anything to do with the great and dreadful humanitarian problems and disasters of our age; and those who are above all concerned with these humanitarian disasters have no time at all for abstruse issues concerning the aims and methods of physics. And so the connection is never made. As it happens, the methodological neurosis of physics, with which we begin in chapter one, is actually a quite simple matter to grasp. It does not require any real knowledge of physics to understand. Indeed, physicists and philosophers of physics are much more likely to find the argument difficult to follow than are non-experts. We non-scientists can stand back and see the whole wood; scientists, trained to think in terms of the current orthodox conception of science, trapped in the thickets of research, will find this much harder to do. In an attempt to take this peculiar circumstance into account, I have arranged the exposition as follows. In chapter one I give a simple exposition of the case for declaring natural science to suffer from what I call "rationalistic neurosis"; this avoids all technicalities, and goes straight to the heart of the matter. In the appendix I tackle a host of more technical objections that may be made to the elementary argument of chapter one. So, those experts in the fields of the philosophy of physics and philosophy of science who find the argument of chapter one naïve and unconvincing, should consult the appendix before tossing the book away in disgust. I write in the hope that there will be a few who will not dismiss out of hand the suggestion that the question of how we are to go about learning how to live in wiser and more civilized ways might have something to learn from scientific learning, and will take the trouble to pursue the line of argument traced out in this book. I write in the hope that these few will grasp just how desperate our situation is, how urgent the need to change the status quo, and will do everything in their power to alert others to the need to heal the methodological sickness from which our institutions and traditions of learning at present suffer. To begin with, we need a campaigning organization, modelled perhaps on "Friends of the Earth", which might be called something like "Friends of Wisdom". (shrink)

Our philosophical understanding of mental illness is being shaped by neuroscience. However, it has the paradoxical effect of igniting two radically opposed groups of philosophical views. On one side, skepticism and denialism assume that, lacking clear biological mechanisms and etiologies for most mental illnesses, we should infer they are constructions best explained by means of social factors. This is strongly associated with medical nihilism: it considers psychiatry more harmful than benign. On the other side of the divide, naturalism and (...) reductionism are on the look for failures in the biological functioning of the organism whenever a genuine mental illness occurs. Psychiatry as currently practiced, accordingly, exhibits the gaps of an ongoing research programme; a yet to be completed neuroscience would link mental illnesses with identifiable biological mechanisms. Both sides of this divide claim to be fostered by scientific discoveries and advances in neuroscience, when taken at face value. Against this background, we argue instead for a modest view. To that end, we draw attention to some nuances in the scientific realism debate. While contending that neuroscientific theories and models aim to provide true representations of their target systems, and can justifiably claim to have attained some, we argue that our confidence should not be placed beforehand in specific features of these scientific representations. Hence, it would be unwarranted to extract morals for psychiatry from posits (or their absence) in neuroscientific explanations of mental illnesses. To illustrate our position, we examine some recent discoveries in neuroscience concerning bipolar disorder. We conclude by linking our topic to a broader issue in the philosophy of medicine: insofar as psychiatry is a biomedical specialty, its classifications of health and disease are guided by pragmatic concerns, as well as by scientific discoveries. (shrink)

Since 2004, it has been mandated by law that all Danish undergraduate university programmes have to include a compulsory course on the philosophy of science for that particular program. At the Faculty of Science and Technology, Aarhus University, the responsibility for designing and running such courses were given to the Centre for Science Studies, where a series of courses were developed aiming at the various bachelor educations of the Faculty. Since 2005, the Centre has been running a dozen different (...) courses ranging from mathematics, computer science, physics, chemistry over medical chemistry, biology, molecular biology to sports science, geology, molecular medicine, nano science, and engineering. -/- We have adopted a teaching philosophy of using historical and contemporary case studies to anchor broader philosophical discussions in the particular subject discipline under consideration. Thus, the courses are tailored to the interests of the students of the particular programme whilst aiming for broader and important philosophical themes as well as addressing the specific mandated requirements to integrate philosophy, some introductory ethics, and some institutional history. These are multiple and diverse purposes which cannot be met except by compromise. -/- In this short presentation, we discuss our ambitions for using case studies to discuss philosophical issues and the relation between the specific philosophical discussions in the disciplines and the broader themes of philosophy of science. We give examples of the cases chosen to discuss various issues of scientific knowledge, the role of experiments, the relations between mathematics and science, and the issues of responsibility and trust in scientific results. Finally, we address the issue of how and why science students can be interested in and benefit from mandatory courses in the philosophy of their subject. (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. J. Ovarian follicular dynamics: a review with emphasis on the bovine species. Part I: Folliculogenesis and pre‐antral follicle development. Reproduction in domestic animals, v. 45, n. 1, p. 171-179, 2010. AERTS, J. M. J.; BOLS, P. E. J. Ovarian follicular dynamics. A review with emphasis on the bovine species. Part II: Antral development, exogenous influence and future prospects. Reproduction in domestic animals, v. 45, n. 1, p. 180-187, 2010. ALBERTINI, David F.; CARABATSOS, Mary Jo. Comparative aspects of meiotic cell cycle control in mammals. Journal of molecular medicine, v. 76, n. 12, p. 795-799, 1998. AUSTIN, Colin Russell; SHORT, R. Reproduction in mammals. Cambridge, 1972. BAKER, T. G. Oogenesis and ovulation. In. Reproduction in Mammals I Germ Cells and Fertilization, p. 29-30, 1972. BEARDEN, Henry Joe et al. Reproducción animal aplicada. México: Manual Moderno, 1982. BIGGERS, John D.; SCHUETZ, Allen W. Oogenesis. University Park Press, 1972. BINELLI, Mario; MURPHY, Bruce D. Coordinated regulation of follicle development by germ and somatic cells. Reproduction, Fertility and Development, v. 22, n. 1, p. 1-12, 2009. CHIARINI-GARCIA, Helio; RUSSELL, Lonnie D. High-resolution light microscopic characterization of mouse spermatogonia. Biology of reproduction, v. 65, n. 4, p. 1170-1178, 2001. CHOUDARY, J. B.; GIER, H. T.; MARION, G. B. Cyclic changes in bovine vesicular follicles. Journal of animal science, v. 27, n. 2, p. 468-471, 1968. CLERMONT, Yves; PEREY, Bernard. Quantitative study of the cell population of the seminiferous tubules in immature rats. American Journal of Anatomy, v. 100, n. 2, p. 241-267, 1957. COSTA, DEILER SAMPAIO; PAULA, T. A. R. Espermatogênese em mamíferos. Scientia, v. 4, 2003. CUNNINGHAM, James. Tratado de fisiologia veterinária. Elsevier Health Sciences, 2011. CUPPS, Perry T. (Ed.). Reproduction in domestic animals. Elsevier, 1991. DA SILVA, Emanuel Isaque Cordeiro. 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. A study of the development of testicular function and an evaluation of testicular biopsy in farm animals. 1971. Tese de Doutorado. Royal Veterinary College (University of London). GALLICANO, G. Ian. Composition, regulation, and function of the cytoskeleton in mammalian eggs and embryos. Front Biosci, v. 6, p. D1089-1108, 2001. GILBERT, Scott F. Biología del desarrollo. Ed. Médica Panamericana, 2005. GNESSI, Lucio; FABBRI, Andrea; SPERA, Giovanni. Gonadal peptides as mediators of development and functional control of the testis: an integrated system with hormones and local environment. Endocrine reviews, v. 18, n. 4, p. 541-609, 1997. HAFEZ, Elsayed Saad Eldin; HAFEZ, Bahaa. Reprodução animal. São Paulo: Manole, 2004. HEDGER, Mark P. Testicular leukocytes: what are they doing?. Reviews of reproduction, v. 2, n. 1, p. 38-47, 1997. HUTSON, James C. Testicular macrophages. In: International review of cytology. Academic Press, 1994. p. 99-143. HYTTEL, P. Gametogênese. In. 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. Archives of medical research, v. 32, n. 6, p. 553-558, 2001. MINTZ, Beatrice et al. Embryological phases of mammalian gametogenesis. Embryological phases of mammalian gametogenesis., v. 56, n. Suppl. 1, p. 31-43, 1960. MORALES, M. E. et al. Gametogénesis. I. Revisión de la literatura, con enfoque en la ovogénesis. Medicina Universitaria, v. 8, n. 32, p. 183-9, 2006. NAKATSUJI, NORIO; CHUMA, SHINICHIRO. Differentiation of mouse primordial germ cells into female or male germ cells. International Journal of Developmental Biology, v. 45, n. 3, p. 541-548, 2002. NILSSON, Eric; PARROTT, Jeff A.; SKINNER, Michael K. Basic fibroblast growth factor induces primordial follicle development and initiates folliculogenesis. Molecular and cellular endocrinology, v. 175, n. 1-2, p. 123-130, 2001. REFERÊNCIAS BIBLIOGRÁFICAS -/- NORRIS, David O.; LOPEZ, Kristin H. The endocrinology of the mammalian ovary. In: Hormones and reproduction of vertebrates. Academic Press, 2011. p. 59-72. PEDERSEN, Torben. 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. Biology of reproduction, v. 66, n. 4, p. 1134-1150, 2002. SCARAMUZZI, R. J.; MARTENSZ, N. D.; VAN LOOK, P. F. A. Ovarian morphology and the concentration of steroids, and of gonadotrophins during the breeding season in ewes actively immunized against oestradiol-17β or oestrone. Reproduction, v. 59, n. 2, p. 303-310, 1980. SEIDEL JR, G. E. et al. Control of folliculogenesis and ovulation in domestic animals: puberal and adult function. In: 9th International Congress on Animal Reproduction and Artificial Insemination, 16th-20th June 1980. II. Round tables. Editorial Garsi., 1980. p. 11-16. SKINNER, Michael K. Cell-cell interactions in the testis. Endocrine Reviews, v. 12, n. 1, p. 45-77, 1991. SMITZ, J. E.; CORTVRINDT, Rita G. The earliest stages of folliculogenesis in vitro. Reproduction, v. 123, n. 2, p. 185-202, 2002. SORENSEN, Anton Marinus. Reproducción animal: principios y prácticas. México, 1982. SUTOVSKY, Peter; MANANDHAR, Gaurishankar. Mammalian spermatogenesis and sperm structure: anatomical and compartmental analysis. In. The sperm cell: Production, maturation, fertilization, regeneration, p. 1-30, 2006. TAZUKE, Salli I. et al. A germline-specific gap junction protein required for survival of differentiating early germ cells. Development, v. 129, n. 10, p. 2529-2539, 2002. VAN STRAATEN, H. W. M.; WENSING, C. J. G. Leydig cell development in the testis of the pig. Biology of Reproduction, v. 18, n. 1, p. 86-93, 1978. TURNBULL, K. E.; BRADEN, A. W. H.; MATTNER, P. E. The pattern of follicular growth and atresia in the ovine ovary. Australian Journal of Biological Sciences, v. 30, n. 3, p. 229-242, 1977. WASSARMAN, Paul M. Gametogenesis. Londres: Academic Press, 2012. WROBEL, K.-H.; SÜß, Franz. Identification and temporospatial distribution of bovine primordial germ cells prior to gonadal sexual differentiation. Anatomy and embryology, v. 197, n. 6, p. 451-467, 1998. REFERÊNCIAS BIBLIOGRÁFICAS -/- ZARCO, L. 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)

Debates about obesity in bioethics tend to unfold in predictable epicycles between individual choices and behaviours and the oppressive socio-economic structures constraining them. Here, we argue that recent work from two cutting-edge research programmes in microbiology and social psychology can advance this conceptual stalemate in the literature. We begin in section 1 by discussing two promising lines of obesity research involving the human microbiome and relationship partners. Then, in section 2, we show how this research has made viable novel (...) strategies for fighting obesity, including microbial therapies and dyad-level interventions. Finally, in section 3, we consider objections to our account and conclude by arguing that attention to the most immediate features of our biological and social environment offers a middle ground solution, while also raising important new issues for bioethicists. (shrink)

Gregor Mendel (1822-1884) was the first to explain that certain 'traits' were inherited in plants from one generation to the next. These would later become known as genes. Frederich Miescher in 1869 analyzed a substance from the nucleus of cells, which he therefore called nuclein. Further study of nuclein revealed that it contained elements like hydrogen, oxygen, nitrogen and phosphorous, with a specific ratio of nitrogen to phosphorous. Then in 1878 Albrecht Kossel determined that nuclein contained nucleic acid, from which (...) he isolated five nucleobases (nitrogen compounds now referred to by the letters C, G, A, T, U representing cytosine, guanine, adenine, thymine, and uracil). It was also discovered that ribose, a sugar was present in the nuclein compound. What Miescher had isolated from the cell nucleus was actually what would latter be identified as DNA (Deoxyribonucleic acid). C. H. Waddington (1905 - 1975) first proposed the term “epigenetics” in 1942 to describe the region between the gene and the whole organism (phenotype) [3]. Today, what is called the epigenome refers to all the chromosomal modifications, DNA modifications, chromatin protein modifications and their complexes. It is the epigenome that determines both the expression of the genes and their inheritance. R. A. Jorgenson reports [4], "Many of these modifications appear to be “programmable” and to be “read out” to influence chromosomal functions." Nobel laureate Barbara McClintock stated this revolutionary proposal more clearly in her Nobel lecture [5], “to determine the extent of knowledge the cell has of itself, and how it utilizes this knowledge in a ‘thoughtful’ manner when challenged.” In 1960 R. A. Brinks suggested that chromosomes possess a paragenetic function in addition to their genetic function [7]. The physical nature of the paragenetic function is characterized by the variety of forms or states of chromatin that can reside at any genetic locus. While the genetic function is stable, the paragenetic function is labile and programmable in ontogeny. It is this latter function that allows organisms to transfer informational macromolecules (RNA and proteins) in a systematic and regulated manner over what is known as the “RNA information superhighway.” Given this capacity, organisms may be able to store information at numerous genetic loci in the form of paragenetic chromatin states, which can be reprogrammed during ontogeny or environmental stress [8]. This reprogrammable system could operate over the whole organism as a storage device, allowing it to make informed ‘decisions' during growth and development, or in response to the environment. Such processing capacity could be considered a form of ‘intelligence,' which also could be passed on to future generations. The study of the flow of information within and between cells and organisms represents the cutting edge of modern biological research. While physical correlates of cognitive behavior in living organisms are being discovered, it does not spell reduction to such correlates. The electronic activity within the physical components of a radio, for example, may be minutely determined, but ultimately it is not merely the electrical activity that produces the intelligent speech that is heard. Only the intelligent person whose voice is being broadcast through the radio can explain that. Without the broadcaster, the radio would sit silently even though fully functional. An organism without its living agency also appears to be devoid of metabolic activity although all the chemical components are fully present. How to connect life to matter will be the ultimate challenge that has to be met. This will prove to be a philosophical problem we hope to address in the near future. (shrink)

All SETI (Search for Extraterrestrial Intelligence) programmes that were conceived and put into practice since the 1960s have been based on anthropocentric ideas concerning the definition of intelligence on a cosmic-wide scale. Brain-based neuronal intelligence, augmented by AI, are currently thought of as being the only form of intelligence that can engage in SETI-type interactions, and this assumption is likely to be connected with the dilemma of the famous Fermi paradox. We argue that high levels of intelligence and cognition (...) inherent in ensembles of bacteria are much more likely to be the dominant form of cosmic intelligence, and the transfer of such intelligence is enabled by the processes of panspermia. We outline the main principles of bacterial intelligence, and how this intelligence may be used by the planetary-scale bacterial system, or the bacteriosphere, through processes of biological tropism, to connect to any extra-terrestrial microbial forms, independently of human interference. (shrink)

In their seminal work, Dialectics of Enlightenment, Horkheimer and Adorno interpreted capitalism as the irrational monetization of nature. In the present work, I analyze three 21st century concepts, Anthropocene, Capitalocene and Machinocene, in light of Horkheimer and Adorno’s arguments and recent arguments from the philosophy of biology. The analysis reveals a remarkable prescience of the term “instrumental reason”, which is present in each of the three concepts in a profound and cryptic way. In my interpretation, the term describes the propensity (...) of science based on the notion of physicalism to interpret nature as the machine analyzable and programmable by the human reason. As a result, the Anthropocene concept is built around the mechanicist model, which may be presented as the metaphor of the car without brakes. In a similar fashion, the Machinocene concept predicts the emergence of the mechanical mind, which will dominate nature in the near future. Finally, the Capitalocene concept turns a perfectly rational ambition to expand knowledge into an irrational obsession with over-knowledge, by employing the institutionalized science as the engine of capitalism without brakes. The common denominator of all three concepts is the irrational propensity to legitimize self-destruction. Potential avenues for countering the effects of “instrumental reason” are suggested. (shrink)

This dissertation attempts to apply Imre Lakatos' Philosophy of Science to the research program of Mary Schweitzer and staff, presented in two papers published in Science: one of them about the discovery of T-Rex with soft tissue and a later one reporting the detection of collagen. This case-study highlights the more general applicability of Lakatos' Methodology of Scientific Research Programmes to scientific practice in Evolutionary Biology.

ABSTRACT of The Flying Termite by L.L. Katona -/- In this book I would like to show the term “intelligence“ has a universal, non-anthropomorphic meaning. We can perceive intelligence in dogs, dolphins or gorillas without understanding of it, but intelligence can be also seen in many other things from insects and the Solar System to elementary particles or the rules of a triangle. But that doesn’t mean intelligence comes from Intelligent Design, yet alone a Designer, they seems to be the (...) “state of things” in the Universe. One of the most known law of the triangle is “the measures of the interior angles of a triangle in Euclidean space always add up to 180 degrees.” The question is this law exists without any existing triangle, or not? If there is an encrypted program in DNA, what determines which living being will be a chameleon or a cat, where this program lies? Whether in the DNA itself? But DNA also programmed. Where is the program of a computer? You may say in the hard drive or in the head of the programmer. But the hard drive or the head of the programmer is just the vehicle of the program; when you dissect a computer or a programmer’s head you will not find the program, just the diodes or cells connected by pack of orders. And you won’t see the orders in them. In my book I want to show the works of this elusive intelligence with some interesting contradictions between causality, determinism, teleology, and dependent formation through biology, mathematics and logic. If I need to summarize my idea I can use the following parable. A priest and an atheist are playing with a beach ball in a pool, in a two-dimensioned plane. They are rod-people with a circle. What they can’t perceive is this plane is a part of a three-dimensioned world, where the wind is blowing, so they can’t understand why the ball disappearing continuously, and why it comes back at another point randomly. Both of them have a hint, the ball is a sphere, and the space is three-dimensioned, but instead of trying to work in team to understand this curious phenomena of the Beach Ball, they’re using the old method: insulting each other. (shrink)

Neo-Darwinism is simply incorrect and it is an indictment of modern biology that the Theory of Evolution has not been updated in the light of research that can easily be conducted by anyone with a programmable computer.

The impressive variation amongst biological individuals generates many complexities in addressing the simple-sounding question what is a biological individual? A distinction between evolutionary and physiological individuals is useful in thinking about biological individuals, as is attention to the kinds of groups, such as superorganisms and species, that have sometimes been thought of as biological individuals. More fully understanding the conceptual space that biological individuals occupy also involves considering a range of other concepts, such as life, (...) reproduction, and agency. There has been a focus in some recent discussions by both philosophers and biologists on how evolutionary individuals are created and regulated, as well as continuing work on the evolution of individuality. (shrink)

Direct neurological and especially imaging-driven investigations into the structures essential to naturally occurring cognitive systems in their development and operation have motivated broadening interest in the potential for artificial consciousness modeled on these systems. This first paper in a series of three begins with a brief review of Boltuc’s (2009) “brain-based” thesis on the prospect of artificial consciousness, focusing on his formulation of h-consciousness. We then explore some of the implications of brain research on the structure of consciousness, finding limitations (...) in biological approaches to the study of consciousness. Looking past these limitations, we introduce research in artificial consciousness designed to test for the emergence of consciousness, a phenomenon beyond the purview of the study of existing biological systems. (shrink)

“Impermanent Biological Phenomena” July 2021, Buddhism and Culture (a Korean-language Buddhist magazine sponsored by the Foundation for the Promotion of Korean Buddhism), Korea.

This third paper locates the synthetic neurorobotics research reviewed in the second paper in terms of themes introduced in the first paper. It begins with biological non-reductionism as understood by Searle. It emphasizes the role of synthetic neurorobotics studies in accessing the dynamic structure essential to consciousness with a focus on system criticality and self, develops a distinction between simulated and formal consciousness based on this emphasis, reviews Tani and colleagues' work in light of this distinction, and ends by (...) forecasting the increasing importance of synthetic neurorobotics studies for cognitive science and philosophy of mind going forward, finally in regards to most- and myth-consciousness. (shrink)

Science and scientific knowledge have been questioned in many ways for a long period of time. Especially, after the scientific revolution of 16th- and 17th-century Europe, science and its knowledge have been mainly accepted one of the most valuable and trustable information. However, in 20th century, autonomy of scientific knowledge and its dominant position over other kinds of knowledge have been mainly criticised. Social and other factors that were tried to be excluded before have been incorporated into the work by (...) the influence of the Strong Programme. In this article, it will be argued that while people are presenting scientific knowledge, their interests, beliefs and the communities they are involved in are also shown to be effective in producing this information. Thus, the desired result is that it is not reasonable to talk about the absolute autonomy of scientific knowledge. (shrink)

1900 was a remarkable year for science. Several ground-breaking events took place, in physics, biology and psychology. Planck introduced the quantum concept, the work of Mendel was rediscovered, and Sigmund Freud published The Interpretation of Dreams . These events heralded the emergence of completely new areas of inquiry, all of which greatly affected the intellectual landscape of the 20 th century, namely quantum physics, genetics and psychoanalysis. What do these developments have in common? Can we discern a family likeness, a (...) basic affinity between them, so that we can use the one to deepen our understanding of the other? One common denominator is that they open up realms of inquiry that are significantly different from the world of everyday experience, namely the realm of elementary particles, of genes and genomes, and of the unconscious. But to what extent can we meaningfully argue, for instance, that the genome is the biological unconscious, and the unconscious the psychic genome? To address these questions, I will build on the work of two key intellectual figures who have explored the affinities of these developments in depth, namely Erwin Schrödinger (a quantum physicist and avid reader of Schopenhauer who initiated molecular biology) and Jacques Lacan (who reframed the specificity of psychoanalysis with the help of 20 th century science: the era of structural linguistics, but also of quantum physics, molecular biology, bioinformatics and DNA). (shrink)

An influential position in the philosophy of biology claims that there are no biological laws, since any apparently biological generalization is either too accidental, fact-like or contingent to be named a law, or is simply reducible to physical laws that regulate electrical and chemical interactions taking place between merely physical systems. In the following I will stress a neglected aspect of the debate that emerges directly from the growing importance of mathematical models of biological phenomena. My main (...) aim is to defend, as well as reinforce, the view that there are indeed laws also in biology, and that their difference in stability, contingency or resilience with respect to physical laws is one of degrees, and not of kind . (shrink)

This selective review explores biologically inspired learning as a model for intelligent robot control and sensing technology on the basis of specific examples. Hebbian synaptic learning is discussed as a functionally relevant model for machine learning and intelligence, as explained on the basis of examples from the highly plastic biological neural networks of invertebrates and vertebrates. Its potential for adaptive learning and control without supervision, the generation of functional complexity, and control architectures based on self-organization is brought forward. Learning (...) without prior knowledge based on excitatory and inhibitory neural mechanisms accounts for the process through which survival-relevant or task-relevant representations are either reinforced or suppressed. The basic mechanisms of unsupervised biological learning drive synaptic plasticity and adaptation for behavioral success in living brains with different levels of complexity. The insights collected here point toward the Hebbian model as a choice solution for “intelligent” robotics and sensor systems. Keywords: Hebbian learning; synaptic plasticity; neural networks; self-organization; brain; reinforcement; sensory processing; robot control . (shrink)

Several authors have used the notion of causal specificity in order to defend non-parity about genetic causes (Waters 2007, Woodward 2010, Weber 2017, forthcoming). Non-parity in this context is the idea that DNA and some other biomolecules that are often described as information-bearers by biologists play a unique role in life processes, an idea that has been challenged by Developmental Systems Theory (e.g., Oyama 2000). Indeed, it has proven to be quite difficult to state clearly what the alleged special role (...) of genetic causes consists in. In this paper, I show that the set of biomolecules that are normally considered to be information-bearers (DNA, mRNA) can be shown to be the most specific causes of protein primary structure, provided that causal specificity is measured over a relevant space of biological possibilities, disregarding physical as well as logically possible states of the causal variables. (shrink)

Biological atomism postulates that all life is composed of elementary and indivisible vital units. The activity of a living organism is thus conceived as the result of the activities and interactions of its elementary constituents, each of which individually already exhibits all the attributes proper to life. This paper surveys some of the key episodes in the history of biological atomism, and situates cell theory within this tradition. The atomistic foundations of cell theory are subsequently dissected and discussed, (...) together with the theory’s conceptual development and eventual consolidation. This paper then examines the major criticisms that have been waged against cell theory, and argues that these too can be interpreted through the prism of biological atomism as attempts to relocate the true biological atom away from the cell to a level of organization above or below it. Overall, biological atomism provides a useful perspective through which to examine the history and philosophy of cell theory, and it also opens up a new way of thinking about the epistemic decomposition of living organisms that significantly departs from the physicochemical reductionism of mechanistic biology. (shrink)

Since Boorse [Philos Sci 44(4):542–573, 1977] published his paper “Health as a theoretical concept” one of the most lively debates within philosophy of medicine has been on the question of whether health and disease are in some sense ‘objective’ and ‘value-free’ or ‘subjective’ and ‘value-laden’. Due to the apparent ‘failure’ of pure naturalist, constructivist, or normativist accounts, much in the recent literature has appealed to more conciliatory approaches or so-called ‘hybrid accounts’ of health and disease. A recent paper by Matthewson (...) and Griffiths [J Med Philos 42(4):447–466, 2017], however, may bear the seeds for the revival of purely naturalist approach to health and disease. In this paper, I defend their idea of Biological Normativity against recent criticism by Schwartz [J Med Philos Forum Bioethics Philos Med 42(4):485–502, 2017] and hope to help it flower into a revival of naturalist approaches in the philosophy of medicine. (shrink)

Although contemporary metaphysics has recently undergone a neo-Aristotelian revival wherein dispositions, or capacities are now commonplace in empirically grounded ontologies, being routinely utilised in theories of causality and modality, a central Aristotelian concept has yet to be given serious attention – the doctrine of hylomorphism. The reason for this is clear: while the Aristotelian ontological distinction between actuality and potentiality has proven to be a fruitful conceptual framework with which to model the operation of the natural world, the distinction between (...) form and matter has yet to similarly earn its keep. In this chapter, I offer a first step toward showing that the hylomorphic framework is up to that task. To do so, I return to the birthplace of that doctrine - the biological realm. Utilising recent advances in developmental biology, I argue that the hylomorphic framework is an empirically adequate and conceptually rich explanatory schema with which to model the nature of organisms. (shrink)

Synthetic biologists aim to generate biological organisms according to rational design principles. Their work may have many beneficial applications, but it also raises potentially serious ethical concerns. In this article, we consider what attention the discipline demands from bioethicists. We argue that the most important issue for ethicists to examine is the risk that knowledge from synthetic biology will be misused, for example, in biological terrorism or warfare. To adequately address this concern, bioethics will need to broaden its (...) scope, contemplating not just the means by which scientific knowledge is produced, but also what kinds of knowledge should be sought and disseminated. (shrink)

The selected effects or ‘etiological’ theory of Proper function is a naturalistic and realist account of biological teleology. It is used to analyse normativity in philosophy of language, philosophy of mind, philosophy of medicine and elsewhere. The theory has been developed with a simple and intuitive view of natural selection. Traits are selected because of their positive effects on the fitness of the organisms that have them. These ‘selected effects’ are the Proper functions of the traits. Proponents argue that (...) this analysis of biological teleology has the unique advantage that the selected effect function of a trait is also a causal explanation of the trait: the trait exists because it performs this function. We show, however, that selected effect functions as currently defined explain the existence of traits only under highly restrictive assumptions about evolutionary dynamics. In many common scenarios in which traits evolve by natural selection, selected effect functions do not explain those traits. This is because definitions of selected effect function extract from any evolutionary scenario only the information that would be explanatorily relevant in the simple evolutionary scenario implicit in those definitions. When applied to more complex scenarios selected effect functions omit the key information that is explanatorily relevant in those scenarios. The assumptions required for selected effect functions to be explanatory are particularly unlikely to hold in the domain that its proponents care most about - the evolution of representation. A more adequate selected effects theory of Proper functions may be possible, but will require much greater attention to the structure of actual evolutionary explanations. (shrink)

García-Barranquero et al explore the desirability of human ageing. They differentiate between chronological and biological views of ageing and contend that the positive aspects of ageing are solely linked to chronological ageing. Consequently, the authors embrace the potential for technological interventions in biological ageing. Contrary to their stance, I argue that there are sometimes desirable aspects associated with biological ageing. Therefore, proposals aiming to eliminate, mitigate or diminish biological ageing are not without problems.

Absolute needs (as against instrumental needs) are independent of the ends, goals and purposes of personal agents. Against the view that the only needs are instrumental needs, David Wiggins and Garrett Thomson have defended absolute needs on the grounds that the verb ‘need’ has instrumental and absolute senses. While remaining neutral about it, this article does not adopt that approach. Instead, it suggests that there are absolute biological needs. The absolute nature of these needs is defended by appeal to: (...) their objectivity (as against mind-dependence); the universality of the phenomenon of needing across the plant and animal kingdoms; the impossibility that biological needs depend wholly upon the exercise of the abilities characteristic of personal agency; the contention that the possession of biological needs is prior to the possession of the abilities characteristic of personal agency. Finally, three philosophical usages of ‘normative’ are distinguished. On two of these, to describe a phenomenon or claim as ‘normative’ is to describe it as value-dependent. A description of a phenomenon or claim as ‘normative’ in the third sense does not entail such value-dependency, though it leaves open the possibility that value depends upon the phenomenon or upon the truth of the claim. It is argued that while survival needs (or claims about them) may well be normative in this third sense, they are normative in neither of the first two. Thus, the idea of absolute need is not inherently normative in either of the first two senses. (shrink)

Is it possible to know anything about life we have not yet encountered? We know of only one example of life: our own. Given this, many scientists are inclined to doubt that any principles of Earth’s biology will generalize to other worlds in which life might exist. Let’s call this the “N = 1 problem.” By comparison, we expect the principles of geometry, mechanics, and chemistry would generalize. Interestingly, each of these has predictable consequences when applied to biology. The surface-to-volume (...) property of geometry, for example, limits the size of unassisted cells in a given medium. This effect is real, precise, universal, and predictive. Furthermore, there are basic problems all life must solve if it is to persist, such as resistance to radiation, faithful inheritance, and energy regulation. If these universal problems have a limited set of possible solutions, some common outcomes must consistently emerge. In this chapter, I discuss the N = 1 problem, its implications, and my response (Mariscal 2014). I hold that our current knowledge of biology can justify believing certain generalizations as holding for life anywhere. Life on Earth may be our only example of life, but this is only a reason to be cautious in our approach to life in the universe, not a reason to give up altogether. In my account, a candidate biological generalization is assessed by the assumptions it makes. A claim is accepted only if its justification includes principles of evolution, but no contingent facts of life on Earth. (shrink)

In this paper I first offer a systematic outline of a series of conceptual novelties in the life-sciences that have favoured, over the last three decades, the emergence of a more social view of biology. I focus in particular on three areas of investigation: (1) technical changes in evolutionary literature that have provoked a rethinking of the possibility of altruism, morality and prosocial behaviours in evolution; (2) changes in neuroscience, from an understanding of the brain as an isolated data processor (...) to the ultrasocial and multiply connected social brain of contemporary neuroscience; and (3) changes in molecular biology, from the view of the gene as an autonomous master of development to the ‘reactive genome’ of the new emerging field of molecular epigenetics. In the second section I reflect on the possible implications for the social sciences of this novel biosocial terrain and argue that the postgenomic language of extended epigenetic inheritance and blurring of the nature/nurture boundaries will be as provocative for neo-Darwinism as it is for the social sciences as we have known them. Signs of a new biosocial language are emerging in several social-science disciplines and this may represent an exciting theoretical novelty for twenty-first social theory. (shrink)

A biologically unavoidable sequence is an infinite gender sequence which occurs in every gendered, infinite genealogical network satisfying certain tame conditions. We show that every eventually periodic sequence is biologically unavoidable (this generalizes König's Lemma), and we exhibit some biologically avoidable sequences. Finally we give an application of unavoidable sequences to cellular automata.

Modern life sciences research is increasingly relying on artificial intelligence (AI) approaches to model biological systems, primarily centered around the use of machine learning (ML) models. Although ML is undeniably useful for identifying patterns in large, complex data sets, its widespread application in biological sciences represents a significant deviation from traditional methods of scientific inquiry. As such, the interplay between these models and scientific understanding in biology is a topic with important implications for the future of scientific research, (...) yet it is a subject that has received little attention. Here, we draw from an epistemological toolkit to contextualize recent applications of ML in biological sciences under modern philosophical theories of understanding, identifying general principles that can guide the design and application of ML systems to model biological phenomena and advance scientific knowledge. We propose that conceptions of scientific understanding as information compression, qualitative intelligibility, and dependency relation modelling provide a useful framework for interpreting ML-mediated understanding of biological systems. Through a detailed analysis of two key application areas of ML in modern biological research – protein structure prediction and single cell RNA-sequencing – we explore how these features have thus far enabled ML systems to advance scientific understanding of their target phenomena, how they may guide the development of future ML models, and the key obstacles that remain in preventing ML from achieving its potential as a tool for biological discovery. Consideration of the epistemological features of ML applications in biology will improve the prospects of these methods to solve important problems and advance scientific understanding of living systems. (shrink)

This study used a partial least squares structural equation modelling (PLS-SEM) to estimate curriculum management's direct and indirect effects on university graduate programmes' viability. The study also examined the role of institutional effectiveness in mediating the nexus between the predictor and response variables. This is a correlational study with a factorial research design. The study's participants comprised 149 higher education administrators (23 Faculty Deans and 126 HODs) from two public universities in Nigeria. A structured questionnaire designed by the researchers (...) was used for data collection. The questionnaire was duly validated with an acceptable scale and item content validity indices. The dimensionality of the instrument was determined using exploratory factor analysis (EFA). Convergent validity was based on Average Variance Extracted (AVE), whereas discriminant validity was based on Fornell-Lacker criteria and the Hetero-Trait Mono-Trait (HTMT) ratio. Acceptable composite reliability estimates of internal consistency were reached for the three sub-scales. Following ethical practices, the questionnaire was physically administered to respondents and retrieved afterwards. Smart PLS (version 3.2.9) and SPSS (version 26.0) programs were used for all the statistical analyses. This study uncovered significant direct and indirect effects of curriculum management on the viability of graduate programmes. Institutional effectiveness significantly impacted graduate programmes’ viability while mediating the nexus between curriculum management and graduate programmes’ viability. Curriculum management and institutional effectiveness jointly explained a significant proportion of graduate programmes’ viability variance. The result of this study proved that graduate programmes’ viability depends, to a great extent, on how much curriculum is managed and how effective institutions are with their services. The result of this study can enable institutions seeking to run viable graduate programmes to re-evaluate their curriculum management practices and the effectiveness of their services. (shrink)

In this paper, I set out to critically review the impact of Popperian falsificationism and Lakatosian methodology of scientific research programmes [MSRP] on economic methodology and philosophy of economics. The central thesis I maintain is that neither Popper's falsificationism nor Lakatos' MSRP offers [philosophy of] economics a proper condition for the reception or rejection of economic theories.

This chapter sets the scene for the subsequent philosophical discussions by surveying a number of biological interventions that have been used, or might in the future be used, for the purposes of crime prevention. These interventions are pharmaceutical interventions intended to suppress libido, treat substance abuse or attention deficit-hyperactivity disorder (ADHD), or modulate serotonin activity; nutritional interventions; and electrical and magnetic brain stimulation. Where applicable, we briefly comment on the historical use of these interventions, and in each case we (...) discuss the evidence that they are effective, or might become so with further refinement. The chapter concludes with a comment on some potentially significant differences between the varieties of intervention that we canvass. (shrink)

Synthetic biology is a field of research that concentrates on the design, construction, and modification of new biomolecular parts and metabolic pathways using engineering techniques and computational models. By employing knowledge of operational pathways from engineering and mathematics such as circuits, oscillators, and digital logic gates, it uses these to understand, model, rewire, and reprogram biological networks and modules. Standard biological parts with known functions are catalogued in a number of registries (e.g. Massachusetts Institute of Technology Registry of (...) Standard Biological Parts). Biological parts can then be selected from the catalogue and assembled in a variety of combinations to construct a system or pathway in a microbe. Through the innovative re-engineering of biological circuits and the optimization of certain metabolic pathways, biological modules can be designed to reprogram organisms to produce products or behaviors. Synthetic biology is what is known as a “platform technology”. That is, it generates highly transferrable theoretical models, engineering principles, and know-how that can be applied to create potential products in a wide variety of industries. Proponents suggest that applications of synthetic biology may be able to provide scientific and engineered solutions to a multitude of worldwide problems from health to energy. Synthetic biology research has already been successful in constructing microbial products which promise to offer cheaper pharmaceuticals such as the antimalarial synthetic drug artemisinin, engineered microbes capable of cleaning up oil spills, and the engineering of biosensors that can detect the presence of high concentrations of arsenic in drinking water. One of the potential benefits of synthetic biology research is in its application to biofuel production. It is this application which is the focus of this entry. The term “biofuel” has referred generally to all liquid fuels that are sourced from plant or plant byproducts and are used for energy necessary for transportation vehicles (Thompson 2012). Biofuels that are produced using synthetic biological techniques re-engineer microbes into biofuel factories are a subset of these. (shrink)

The theory of evolution, which provides the conceptual framework for all modern research in organismal biology and informs research in molecular bi- ology, has gone through several stages of expansion and refinement. Darwin and Wallace (1858) of course proposed the original idea, centering on the twin concepts of natural selection and common descent. Shortly thereafter, Wallace and August Weismann worked toward the complete elimination of any Lamarckian vestiges from the theory, leaning in particular on Weismann’s (1893) concept of the separation (...) of soma and germ, resulting in what is some- times referred to as “neo-Darwinism”. (shrink)

We address Steel’s Programme to identify a ‘preferred’ universe of set theory and the best axioms extending ZFC by using his multiverse axioms MV and the ‘core hypothesis’. In the first part, we examine the evidential framework for MV, in particular the use of large cardinals and of ‘worlds’ obtained through forcing to ‘represent’ alternative extensions of ZFC. In the second part, we address the existence and the possible features of the core of MV_T (where T is ZFC+Large Cardinals). In (...) the last part, we discuss the hypothesis that the core is Ultimate-L, and examine whether and how, based on this fact, the Core Universist can justify V=Ultimate-L as the best (and ultimate) extension of ZFC. To this end, we take into account several strategies, and assess their prospects in the light of MV’s evidential framework. (shrink)