Proximate and ultimate causes in evolutionary biology have come to conflate two distinctions. The first is a distinction between immediate and historical causes. The second is between explanations of mechanism and adaptive function. Mayr emphasized the first distinction but many evolutionary biologists use proximate and ultimate causes to refer to the second. I recommend that ‘ultimate cause’ be abandoned as ambiguous.

While admirable, Guala's discussion of reciprocity suffers from a confusion between proximate causes (psychological mechanisms triggering behaviour) and ultimate causes (evolved function of those psychological mechanisms). Because much work on commits this error, I clarify the difference between proximate and ultimate causes of cooperation and punishment. I also caution against hasty rejections of of experimental evidence.

Within evolutionary biology a distinction is frequently made between proximate and ultimate causes. One apparently plausible interpretation of this dichotomy is that proximate causes concern processes occurring during the life of an organism while ultimate causes refer to those processes (particularly natural selection) that shaped its genome. But ultimate causes are not sought through historical investigations of an organisms lineage. Rather, explanations referring to ultimate causes typically emerge from functional (...) analyses. But these functional analyses do not identify causes of any kind, much less ultimate ones. So-called ultimate explanations are not about causes in any sense resembling those of proximate explanations. The attitude, implicit in the term ultimate cause, that these functional analyses are somehow superordinate to those involving proximate causes is unfounded. Ultimate causes are neither ultimate nor causes. (shrink)

It has been over 60 years since Ernst Mayr famously argued for the distinction between proximate and ultimate causes in biology. In the following decades, Mayr’s proximate-ultimate distinction was well received within evolutionary biology and widely regarded as a major contribution to the philosophy of biology. Despite its enormous influence, there has been a persistent controversy on the distinction. It has been argued that the distinction is untenable. In addition, there have been complaints about the (...) pragmatic value of the distinction in biological research. Some even suggest that the distinction should better be abandoned. In contrast, Mayr had consistently maintained the significance of the proximate-ultimate distinction in biology. There are also other attempts to defend the distinction. The paper examines the debate by taking an integrated History and Philosophy of Science (HPS) approach and argues for a functional approach to causal concepts in scientific practice. (shrink)

Romantic love is a phenomenon of immense interest to the general public as well as to scholars in several disciplines. It is known to be present in almost all human societies and has been studied from a number of perspectives. In this integrative review, we bring together what is known about romantic love using Tinbergen’s “four questions” framework originating from evolutionary biology. Under the first question, related to mechanisms, we show that it is caused by social, psychological mate choice, genetic, (...) neural, and endocrine mechanisms. The mechanisms regulating psychopathology, cognitive biases, and animal models provide further insights into the mechanisms that regulate romantic love. Under the second question, related to development, we show that romantic love exists across the human lifespan in both sexes. We summarize what is known about its development and the internal and external factors that influence it. We consider cross-cultural perspectives and raise the issue of evolutionary mismatch. Under the third question, related to function, we discuss the fitness-relevant benefits and costs of romantic love with reference to mate choice, courtship, sex, and pair-bonding. We outline three possible selective pressures and contend that romantic love is a suite of adaptions and by-products. Under the fourth question, related to phylogeny, we summarize theories of romantic love’s evolutionary history and show that romantic love probably evolved in concert with pair-bonds in our recent ancestors. We describe the mammalian antecedents to romantic love and the contribution of genes and culture to the expression of modern romantic love. We advance four potential scenarios for the evolution of romantic love. We conclude by summarizing what Tinbergen’s four questions tell us, highlighting outstanding questions as avenues of potential future research, and suggesting a novel ethologically informed working definition to accommodate the multi-faceted understanding of romantic love advanced in this review. (shrink)

Made famous by Ernst Mayr (1961), the distinction between proximate and ultimate causation in biological explanation is widely seen as a key tenet of evolutionary theory and a central organizing principle for evolutionary research. The study of immediate, individual-level mechanistic causes of development or physiology (“proximate causation”) is distinguished from the study of historical, population-level statistical causes in evolutionary biology (“ultimate causation”). Since evolutionary developmental biology (evo-devo) is a field that explicitly uses so-called “ (...) class='Hi'>proximate” sciences such as developmental biology, morphology, and embryology in the study of evolution, it challenges the standard construal of the proximate-ultimate distinction and its associated account of causation. The exact nature of the challenge and its ramifications for the viability of the distinction more broadly are contested, but these conceptual questions are central to the status and significance of evo-devo in contemporary evolutionary biology. (shrink)

The validity and utility of the proximate-ultimate distinction in biology have recently been under debate. Opponents of the distinction argue that it rules out individual-level organismic processes from evolutionary explanations, thereby leading to an unfounded separation between organismic causation and evolutionary causation. Proponents of the proximate-ultimate distinction, on the other hand, argue that it serves an important epistemological role in forming different kinds of explanation-seeking questions in biology. In this paper we offer an interpretation the (...) class='Hi'>proximate-ultimate distinction not only as a means of forming explanation-seeking questions, but also as a distinction that can help highlight the way in which individual-level organismic processes can be evolutionary causes. We do this by interpreting the distinction between proximate and ultimate causes as a distinction between structuring and triggering causes. (shrink)

In 1961, Ernst Mayr published a highly influential article on the nature of causation in biology, in which he distinguished between proximate and ultimate causes. Mayr argued that proximate causes (e.g. physiological factors) and ultimate causes (e.g. natural selection) addressed distinct ‘how’ and ‘why’ questions and were not competing alternatives. That distinction retains explanatory value today. However, the adoption of Mayr’s heuristic led to the widespread belief that ontogenetic processes are irrelevant to evolutionary (...) questions, a belief that has (1) hindered progress within evolutionary biology, (2) forged divisions between evolutionary biology and adjacent disciplines and (3) obstructed several contemporary debates in biology. Here we expand on our earlier (Laland et al. in Science 334:1512–1516, 2011) argument that Mayr’s dichotomous formulation has now run its useful course, and that evolutionary biology would be better served by a concept of reciprocal causation, in which causation is perceived to cycle through biological systems recursively. We further suggest that a newer evolutionary synthesis is unlikely to emerge without this change in thinking about causation. (shrink)

Strong reciprocity has recently been subject to heated debate. In this debate, the “West camp” :231–262, 2011), which is critical of the case for SR, and the “Laland camp” :1512–1516, 2011, Biol Philos 28:719–745, 2013), which is sympathetic to the case of SR, seem to take diametrically opposed positions. The West camp criticizes advocates of SR for conflating proximate and ultimate causation. SR is said to be a proximate mechanism that is put forward by its advocates as (...) an ultimate explanation of human cooperation. The West camp thus accuses advocates of SR for not heeding Mayr’s original distinction between ultimate and proximate causation. The Laland camp praises advocates of SR for revising Mayr’s distinction. Advocates of SR are said to replace Mayr’s uni-directional view on the relation between ultimate and proximate causes by the bi-directional one of reciprocal causation. The paper argues that both the West camp and the Laland camp misrepresent what advocates of SR are up to. The West camp is right that SR is a proximate cause of human cooperation. But rather than putting forward SR as an ultimate explanation, as the West camp argues, advocates of SR believe that SR itself is in need of ultimate explanation. Advocates of SR tend to take gene-culture co-evolutionary theory as the correct meta-theoretical framework for advancing ultimate explanations of SR. Appearances notwithstanding, gene-culture coevolutionary theory does not imply Laland et al.’s notion of reciprocal causation. “Reciprocal causation” suggests that proximate and ultimate causes interact simultaneously, while advocates of SR assume that they interact sequentially. I end by arguing that the best way to understand the debate is by disambiguating Mayr’s ultimate-proximate distinction. I propose to reserve “ultimate” and “proximate” for different sorts of explanations, and to use other terms for distinguishing different kinds of causes and different parts of the total causal chain producing behavior. (shrink)

Laland and colleagues have sought to challenge the proximate–ultimate distinction claiming that it imposes a unidirectional model of causation, is limited in its capacity to account for complex biological phenomena, and hinders progress in biology. In this article the core of their argument is critically analyzed. It is claimed that contrary to their claims Laland et al. rely upon the proximate–ultimate distinction to make their points and that their alternative conception of reciprocal causation refers to phenomena (...) that were already accounted for by standard theory. (shrink)

It's been 41 years since the publication of Ernst Mayr's Cause and Effect in Biology wherein Mayr most clearly develops his version of the influential distinction between ultimate and proximate causes in biology. In critically assessing Mayr's essay I uncover false statements and red-herrings about biological explanation. Nevertheless, I argue to uphold an analogue of the ultimate/proximate distinction as it refers to two different kinds of explanations, one dynamical the other statistical.

In this dissertation I address three philosophical problems in the philosophy of biology united by the underlying, and interlocking, issues of the explanatory role of teleological, normative, and representational concepts in biology. In the first chapter, I argue that extant accounts of functions have foundered on a problem I dub the Dysfunction Dilemma, and I offer a way to move forward. Functions are of philosophical interest because the concept plays an important explanatory role in biology, and other sciences, but is (...) apparently teleological and normative –an item’s function is its end, purpose, or goal, and a dysfunctional item is one that fails to realize a particular norm, performance of its proper function. Extant accounts of function do not successfully capture both its explanatory import and its normative and teleological character, either painting dysfunction as conceptually incoherent or painting functions as epiphenomenal. I diagnose the source of the dilemma and offer an account of functional explanation which avoids it. In the second chapter, I defend the idea that genes, like words or concepts, have intentionality or representational/semantic content. First, I argue that while the standard motivation for this idea is mechanistic analogies between molecular mechanisms and digital computation, Ernst Mayr, one of the architects of the modern synthesis in biology, has provided an alternative and more defensible justification for taking the idea seriously. Parallel to the standard case for representational and computational theories of cognition, Mayr’s justification rests on the need to invoke representations to explain end-directed processes. I then put Mayr’s argument to work answering three kinds of objection to the idea that genes are representational: 1) Genetic representations are causally or explanatorily inert, 2) Genetic processes are too complex and context sensitive to be captured by representational concepts, and 3) Details of molecular biology undermine representational conceptions of genes. In the final chapter, I develop a historical and philosophical interpretation of Ernst Mayr’s distinction between proximate and ultimate causes and its relation to Niko Tinbergen’s "four questions." Both frameworks have been influential in the biological study of behavior and are typically presented as a single framework, with Mayr’s proximate-ultimate distinction corresponding to one of two axes dividing Tinbergen’s four questions. I argue that this interpretation misrepresents Mayr’s and Tinbergen’s philosophical positions. I offer a positive account of both frameworks and their relation to one another, and I argue that contrary to many critics, Mayr’s proximate-ultimate distinction, when properly interpreted, does not present a barrier to the project of integrating biological work on Tinbergen’s four questions. (shrink)

In response to recent work on the nature of human aggression, and to shed light on the proximate, as opposed to ultimate, causes of tribal warfare, we present a record of events leading to a fatal Waorani raid on a family from another tribe, followed by a detailed first-person observation of the behavior of the raiders as they prepared themselves for war, and upon their return. We contrast this attack with other Waorani aggressions and speculate on evidence (...) regarding their hormonal underpinnings. On-the-ground ethnographic observations are examined in light of the neuroendocrinological literature. The evidence suggests a chain of causality in launching lethal violence, beginning with a perceived injury, culminating in a massacre, and terminating in rejoicing. Although no blood or saliva samples were taken at the time of this raid, the behaviors were consistent with a deliberate initiation of the hormonal cascade characterizing the “fight-or-flight” response, along with other hormonal changes. We conclude with observations on the stratified interrelationships of the cognitive, social, emotional, and neuroendocrinological causes of aggression leading to coalitional male homicide. (shrink)

Ernst Mayr''s distinction between ultimate and proximate causes is justly considered a major contribution to philosophy of biology. But how did Mayr come to this philosophical distinction, and what role did it play in his earlier scientific work? I address these issues by dividing Mayr''s work into three careers or phases: 1) Mayr the naturalist/researcher, 2) Mayr the representative of and spokesman for evolutionary biology and systematics, and more recently 3) Mayr the historian and philosopher of biology. (...) If we want to understand the role of the proximate/ultimate distinction in Mayr''s more recent career as a philosopher and historian, then it helps to consider hisearlier use of the distinction, in the course of his research, and in his promotion of the professions of evolutionary biology and systematics. I believe that this approach would also shed light on some other important philosophical positions that Mayr has defended, including the distinction between essentialism: and population thinking. (shrink)

Abstract: A common explanatory error concerns conflating epistemological roles between two domains. Here I deal with a special case: when explanations of development replace evolutionary explanations or vice versa. Ernst Mayr famously distinguished between proximate and ultimate causal explanations in biology. His view was central to keeping development outside the theory of evolution. Nevertheless, the explanatory role of developmental processes in evolution is a central theme in current theoretical biology, which has led to several revisions of Mayr’s distinction. (...) In the following, I will review these reviewers to determine whether the integration of development and evolution is based on an appropriate reinterpretation of Mayr’s distinction. The revisionists often propose an interactionist alternative in which proximate and ultimate causes interact to produce evolved traits. The most common interactionist account relies on the idea of reciprocal causation. I will argue that this perspective is still problematic and that the boundaries between explanatory domains are crossed. Instead, we should rethink Mayr’s distinction by adopting an alternative view of evolutionary causation, the so-called Statisticalist view on natural selection, which claims that the only level of causation is the proximate level. By excluding two different levels of causation, this framework is able to avoid fallacious explanations and fully consider reciprocal causation as a proximate phenomenon. I introduce the concept of “statistical reciprocity” to explain the statistical effects of (proximate) reciprocal causation in ultimate explanations and outline some ideas of “population ontogenetics” as a prominent framework for unifying development and evolution beyond interactionist positions. (shrink)

I introduce a range of examples of different causal hypotheses about human mate selection. The hypotheses I focus on come from evolutionary psychology, fluctuating asymmetry research and chemical signaling research. I argue that a major obstacle facing an integrated biology of human behavior is the lack of a causal framework that shows how multiple proximate causal mechanisms can act together to produce components of our behavior.

The so-called Evolutionary Debunking Arguments are arguments that appeal to the evolutionary genealogy of our beliefs to undermine their justification. When applied to morality, such arguments are intended to undermine moral realism. In this paper I will discuss Andreas Mogensen’s recent effort to secure moral realism against EDAs. Mogensen attempts to undermine the challenge provided by EDAs in metaethics through the distinction between proximate and ultimate causes in biology. The problem with this move is that the (...) class='Hi'>proximate/ultimate distinction is misconceived. If ultimate and proximate causes are properly understood to be complementary, such distinction cannot affect EDAs in metaethics. Therefore, I will argue, Mogensen’s argument fails and moral realism is still in danger. (shrink)

What motivates grandparents to their altruism? We review answers from evolutionary theory, sociology, and economics. Sometimes in direct conflict with each other, these accounts of grandparental investment exist side-by-side, with little or no theoretical integration. They all account for some of the data, and none account for all of it. We call for a more comprehensive theoretical framework of grandparental investment that addresses its proximate and ultimate causes, and its variability due to lineage, values, norms, institutions (e.g., (...) inheritance laws), and social welfare regimes. This framework needs to take into account that the demographic shift to low fecundity and mortality in economically developed countries has profoundly altered basic parameters of grandparental investment. We then turn to the possible impact of grandparental acts of altruism, and examine whether benefits of grandparental care in industrialized societies may manifest in terms of less tangible dimensions, such as the grandchildren's cognitive and verbal ability, mental health, and well-being. Although grandparents in industrialized societies continue to invest substantial amounts of time and money in their grandchildren, we find a paucity of studies investigating the influence that this investment has on grandchildren in low-risk family contexts. Under circumstances of duress there is converging evidence that grandparents can provide support that helps to safeguard their children and grandchildren against adverse risks. We conclude by discussing the role that grandparents could play in what has been referred to as Europe's demographic suicide. (shrink)

Research on animal behavior is typically organized according to a combination of two influential frameworks: Ernst Mayr’s distinction between proximate and ultimate causes, and Niko Tinbergen’s “four questions”. My aim is to debunk two common interpretive misconceptions about Mayr’s proximate–ultimate distinction and its relationship to Tinbergen’s four questions, and to offer a new interpretation that avoids both. The first misconception is that the proximate–ultimate distinction maps cleanly onto Tinbergen’s four questions, marking a boundary (...) between Tinbergen’s evolutionary and survival value questions versus developmental and mechanistic questions. The second is that Mayr’s proximate–ultimate distinction is meant to rule out the relevance of proximate causes to evolutionary explanations. I argue that neither is plausible given the text and Mayr’s philosophical aims, namely, to argue that evolutionary biology cannot be reduced to either the physical sciences or to other areas of biology. Through a reconstruction of Mayr’s anti-reductionist argument, I develop an interpretation according to which the proximate–ultimate distinction marks two ways that teleological reasoning can be naturalistically grounded in biology, corresponding to Mayr’s distinction between teleonomic and adapted systems. Mayr distinguishes reduction, which the proximate–ultimate distinction is meant to block, from analysis, through which he allows that proximate causes, causes that are neither proximate nor ultimate, and chance can all contribute to evolutionary explanations. I conclude by suggesting some ways in which the interpretation defended here reframes our understanding of Mayr’s disagreements with some evolutionary-developmental biologists. (shrink)

I argue here that Christian ethical responses to Artificial Intelligence (AI) ought to take on, largely, two different approaches. The first considers proximate ethical concerns related to AI. This ethical approach most often considers more immediate personal and socio-political repercussions and the kind of impact that is occurring now or in the very near future. Proximate ethics of this type includes discussion about fairness, accountability, sustainability and transparency. The second concerns ultimate ethics which focuses on the longer-term (...) impact and implications of AI. Examples of this type might include issues of uniqueness, deep societal transformation and inequality, changes to personal character and even the role AI might have in God's ultimate economy of creation and grace. My contention is that the Christian church needs to attend to both approaches to AI and that when it focuses too myopically on one at the expense of the other it often eclipses the entire witness of the church in our technological society. (shrink)

Ernst Mayr’s classical work on the nature of causation in biology has had a huge influence on biologists as well as philosophers. Although his distinction between proximate and ultimate causation recently came under criticism from those who emphasize the role of development in evolutionary processes, the formal relationship between these two notions remains elusive. Using causal graph theory, this paper offers a unified framework to systematically translate a given “proximate” causal structure into an “ultimate” evolutionary response, (...) and illustrates evolutionary implications of various kinds of causal mechanisms including epigenetic inheritance, maternal effects, and niche construction. These results not only reveal the essential interplay between proximate and ultimate causation in the study of evolution, but also provide a formal method to evaluate or discover non-standard or yet unknown evolutionary phenomena. (shrink)

The human faculty of moral judgment is not well suited to address problems, like climate change, that are global in scope and remote in time. Advocates of ‘moral bioenhancement’ have proposed that we should investigate the use of medical technologies to make human beings more trusting and altruistic, and hence more willing to cooperate in efforts to mitigate the impacts of climate change. We survey recent accounts of the proximate and ultimate causes of human cooperation in order (...) to assess the prospects for bioenhancement. We identify a number of issues that are likely to be significant obstacles to effective bioenhancement, as well as areas for future research. (shrink)

Consciousness, as a higher-order cognitive capacity allowing for the explicit representation of abstract mental states, might be the incidental byproduct of design features from other adaptive systems, such as those governing expansion of the frontal lobes in primates. Although such abilities may have occurred entirely by chance, the standardized entrenchment of this representational capacity in human cognition may have posed engineering dilemmas for natural selection in that consciousness could not be easily removed without disrupting the adaptive features of other design (...) solutions. If so, then those organisms saddled with the burden of higher-order representation by the occurrence of these chance events were suddenly assaulted with a series of social problems previously unencountered by any other species in evolutionary history. Such consciousness-based problems constituted enormous selective pressure for generating ancestrally adaptive psychological programs designed to cope with them. Each of these design solutions was, by necessity, generated and progressively pruned over an extraordinarily short span of geological time. In addition, these programs ran into conflict with more ancient primate social adaptations — such as those underlying sexual coercion and violence — that did not evolve to be sensitive to the epistemic positions of others. These mosaic processes have likely resulted in selection for innumerable algorithmic properties driving human-specific behaviors which are both proximally and ultimately caused by consciousness. Consciousness by itself should be classified as maladaptive; what is adaptive are those psychological programs in place to support its incidental and problematic arrival in the human brain. (shrink)

A common explanatory error in science concerns the conflation of the epistemological roles between two domains. Here we will address a specific case: when explanations of development replace evolutionary explanations or vice versa. Ernst Mayr famously distinguished between proximate and ultimate causal explanations in biology. His view was central to the Modern Synthesis’ exclusion of development from evolutionary theory. Nonetheless, the explanatory role of developmental processes in evolution is a central theme in current theoretical biology which has prompted (...) several revisions of Mayr’s distinction. Here we will review these reviewers to determine whether the integration of development and evolution is based on an appropriate reinterpretation of Mayr’s distinction. In many cases, revisionists suggest an interactionist alternative, in which proximate and ultimate causes interact to produce evolved traits. The most frequent interactionist account relies on the idea of reciprocal causation. We will argue that this perspective is still problematic and that the boundaries between explanatory domains are crossed. Instead, we should rethink Mayr’s distinction by adopting an alternative view of evolutionary causation, the so-called Statisticalist view, which maintains that the only level of causation is the individual level. By ruling out two different levels of causation, this framework is appropriate to avoid fallacious explanations and reconsider reciprocal causation entirely as a proximate phenomenon. We introduce the concept of “statistical reciprocity” to explain the statistical effects of reciprocal causality in populations and outline some ideas of “population ontogenetics” as a prominent framework for unifying development and evolution beyond interactionist positions. (shrink)

ABSTRACT: A systematic reconstruction of Chrysippus’ theory of causes, grounded on the Stoic tenets that causes are bodies, that they are relative, and that all causation can ultimately be traced back to the one ‘active principle’ which pervades all things. I argue that Chrysippus neither developed a finished taxonomy of causes, nor intended to do so, and that he did not have a set of technical terms for mutually exclusive classes of causes. Rather, the various adjectives (...) which he used for causes had the function of describing or explaining particular features of certain causes in particular philosophical contexts. I challenge the sometimes assumed close connection of Chrysippus’ notion of causation with explanation. I show that the standard view that the distinction between proximate and auxiliary causes and perfect and principal causes corresponds to the distinction between internal and external determining factors is not born out by the evidence, and argue that causes of the two types were not thought to co-operate, but rather conceived of as alternatives. (shrink)

A biological function is supposed to be performed adequately, and hence may fail to do so: this is dysfunction. This raises two questions. One is how to make explicit the way in which function can be discriminated from dysfunction without confusing dysfunction with non-function. The second question is how what is “right” and “wrong” can be legitimated by natural regulatory norms. A function can be viewed as a quality to which at least one variable with a definite set of values (...) is associated. Accordingly, function and dysfunction are the same quality of the same trait, but differ in the values associated to this variable. Dysfunction occurs when the associated set of values does not match this normal state. Biological systems have the epistemic singularity that their existence is the consequence of two distinct causal regimes, the so-called “proximate” and “ultimate” causes, whose convergence defines a system’s prescriptive normality. Each cause imposes restrictive rules that limit the possible ways these systems can putatively exist. When a system is insensitive to ultimate cause, it is determined by proximate causation alone, and hence escapes its own prescriptive norms. In conclusion: The normality of a biological system is defined by the convergence of proximate and ultimate causes. A function is a variable-associated quality whose values are defined relative to the norms of the system. Dysfunction occurs when the set of values of the same variable does not match the normality of the system, which can occur when the system is insensitive to ultimate causation. (shrink)

Arbuscular mycorrhiza (AM), a type of plant‐fungal endosymbiosis, and nodulation, a bacterial‐plant endosymbiosis, are the most ubiquitous symbioses on earth. Recent findings have established part of a shared genetic basis underlying these interactions. Here, we approach root endosymbioses through the lens of the homology and modularity concepts aiming at further clarifying the proximate and ultimate causes for the establishment of these biological systems. We review the genetics that underlie interspecific signaling and its concomitant shift in genetic programs (...) for either partner. Also, through the comparative analysis of genetic modules shared by AM and nodulation symbioses, we identify fundamental nodes in these networks, suggesting the elemental steps that may have permitted symbiotic adaptation. Here, we show that this approach, allied to recent technical advances in the study of genetic systems architecture, can provide clear testable hypotheses for the advancement of our understanding on the evolution and development of symbiotic systems. (shrink)

Both Aristotle and Ernst Mayr present theories of dual explanation in biology, with proximal, clearly physical explanations and more distal, biology-specific explanations. Aristotle’s presentation of final cause explanations in Posterior Analytics relates final causes to the necessary material, formal, and efficient causes that mediate them. Johnson and Leunissen demonstrate the problematic nature of historical and recent interpretations and open the door for a new interpretation consistent with modern evolutionary theory. Mayr’s differentiation of proximate and ultimate/evolutionary (...) class='Hi'>causes provides a key to appropriating Aristotle for modern use, if care is taken to differentiate Mayr’s proximate/distal and mechanical/evolutionary distinctions from his intentional/non-intentional distinction. Building on Mayr, and noting his appropriation of ancient and recent commentary, a strong case can be made for nested explanation, wherein evolutionary explanations are instantiated in systems of mechanical explanations. Biological concepts of organism, gene, function, etc., are etiologically prior to mechanical explanations of function, but temporally posterior to mechanical explanations of historical adaptation. Such an analysis sheds light on recent arguments within evolutionary biology while highlighting the importance of epistemology in contemporary science. (shrink)

This paper applies philosophical tools from the causalists/statisticalists debate to the evo-devo idea of variational tendencies as propensities biasing phenotypic change. It contends that variational properties are present in a statistical sense in some population dynamics models, particularly quantitative genetics ones, providing ultimate variational explanations. It further argues that these properties, contrary to some recent views, cannot be subsumed under natural selection. Finally, it advocates for a causalist interpretation of these explanations, where variational statistical properties indirectly refer to evo-devo’s (...) variational propensities. (shrink)

In most social species, position in the male social hierarchy and reproductive success are positively correlated; in humans, however, this relationship is less clear, with studies of traditional societies yielding mixed results. In the most economically advanced human populations, the adaptiveness of status vanishes altogether; social status and fertility are uncorrelated. These findings have been interpreted to suggest that evolutionary principles may not be appropriate for the explanation of human behavior, especially in modern environments. The present study tests the adaptiveness (...) of social status with actual mating and reproductive data in a representative sample of males from an industrial society. Reproductive success, even when assessed by a more reliable measure ofactualmale fertility than the one commonly used, fails to correlate with social status. In striking contrast, however, status is found to be highly correlated withpotentialfertility, as estimated from copulation frequency. Status thus accounts for as much as 62% of the variance in thisproximatecomponent of fitness. This pattern is remarkably similar to what is found in many traditional societies and would result in a substantial positive relationship between cultural and reproductive success in industrial populations were it not for the novel conditions imposed by contraception and monogamy. Various underlying mechanisms are suggested for these findings, illustrating the value of current behavioral and reproductive data in the study of adaptation. It is concluded that evolutionary explanations of human behavior remain entirely relevant in modern societies. (shrink)

Transcending reviewed proximate theories, Van Lange et al.'s CLASH model attempts to ultimately explain the poleward declension of aggression and violence. Seasonal cold is causal, but, we contend, principally as an ecologically relevant evolutionary pressure. We further argue that futurity and restraint are life history variables, and that Life History Theory evolutionarily explains the biogeography of aggression and violence as strategic adaptation.

Ernst Mayr proposed a distinction between “proximate”, mechanistic, and “ultimate”, evolutionary, causes of biological phenomena. This dichotomy has influenced the thinking of many biologists, but it is increasingly perceived as impeding modern studies of evolutionary processes, including study of “niche construction” in which organisms alter their environments in ways supportive of their evolutionary success. Some still find value for this dichotomy in its separation of answers to “how?” versus “why?”questions about evolution. But “why is A?” questions about (...) evolution necessarily take the form “how does A occur?”, so this separation is illusory. Moreover, the dichotomy distorts our view of evolutionary causality, in that, contra Mayr, the action of natural selection, driven by genotype-phenotype-environment interactions which constitute adaptations, is no less “proximate” than the biological mechanisms which are altered by naturally selected genetic variants. Mayr’s dichotomy thus needs replacement by more realistic, mechanistic views of evolution. From a mechanistic viewpoint, there is a continuum of adaptations from those evolving as responses to unchanging environmental pressures to those evolving as the capacity for niche construction, and intermediate stages of this can be identified. Some biologists postulate an association of “phenotypic plasticity” (phenotype-environment covariation with genotype held constant) with capacity for niche construction. Both “plasticity” and niche construction comprise wide ranges of adaptive mechanisms, often fully heritable and resulting from case-specific evolution. Association of “plasticity” with niche construction is most likely to arise in systems wherein capacity for complex learning and behavioral flexibility have already evolved. (shrink)

Models of the architecture of mature cognitive systems can inform the study of normal and disordered cognitive development, if one distinguishes between proximal and distal causes of performance. The assumption of residual normality need not be made in order to apply adult models to performance early in development, because these models can be modified to reflect the results of compensatory processing.

Evolutionary explanations of behavior are special in that they involve both proximate and ultimate components. Proximately, evolutionary accounts posit mechanisms that generate observed patterns of behavior. At the ultimate level, evolutionary accounts explain the existence of these proximate mechanisms via evolutionary processes such as selection or drift acting in the past. Does positing or accepting such explanations carry any risks? Here I consider two kinds of risk, epistemic and ethical. Epistemic risk is the risk of being (...) wrong about a matter of fact, such as whether a posited empirical pattern actually exists, or that a particular set of causes explains it. Ethical risk is the risk of causing harm. If the acceptance of a particular explanation for an observed pattern of behavior entails real-world consequences that could lead to harm, then epistemic risk entails ethical risk. Here I consider these risks in cases of evolutionary explanation, and ask whether evolutionary explanations might carry special risks that we should consider when developing and testing evolutionary hypotheses. (shrink)

Thomas Aquinas studied Aristotle at the newly founded University of Naples and, as a Dominican friar, at Paris and at Cologne under Albert the Great. In addition he was well acquainted with the doctrine of the Fathers of the Church, especially Augustine, who shows elements of Neoplatonism. This dual influence upon Aquinas's thinking is apparent in his doctrine on the presence of the natural elements in a mixed body. He asserts that the elements remain virtually in the mixture by the (...) preservation of their powers of acting . This is possible, he explains, because the proper accidents of an element are caused by the elemental substantial form as formal cause, as final cause, and "in a way" as agent cause. Since efficient causality in the proper sense implies that one thing act on another as other, the substantial form of the element is said to be an efficient cause by emanation of the proper accidents . ;The proper operations of a natural substance follow upon its proper accidents, which flow spontaneously from its substantial form. Every natural substance thus exhibits a certain dynamism: it is able to perform its proper operations by reason of the accidents that emanate from its substantial form. Natural substances are secondary and proximate causes. The heavenly bodies, on the other hand, are higher causes moved by the separated substances and, ultimately, by God, the first Cause. Hence, the dynamism of natural substance, although truly present in them, derives from God by way of the heavenly bodies as effects derive from an artisan through his instruments. ;This dissertation examines Aquinas's doctrine as outlined above. The doctrine is first detailed by him and is adopted by the Thomistic schools that followed, becoming the leading doctrine on mixture until the rise of modern science. Chapter One examines the concepts of nature and of element, Chapter Two those of natural substance and of mixture. Chapter Three examines the relationship of substantial form and proper accident in both substantial and accidental changes, with emphasis on the downward motion of heavy bodies. Chapter Four is a close analysis of Aquinas's texts on mixture. This study employs two recently published research tools: the Leonine Commission's edition of the critical texts of Aquinas's philosophical opuscula , and the 49-volume Index Thomisticus. (shrink)

Rice crop diversity hasdecreased dramatically in the recent past.Understanding the causes that underlie theevident genetic erosion is critical for thefood security of subsistence rice farmers andbiodiversity. Our study shows that farmers inthe northeastern Philippines had a markedreduction in rice diversity from 1996 to 1998.The ultimate causes were a drought resultingfrom the El Niño phenomenon in 1997 andflooding due to two successive typhoons in1998. The proximate causes, however, includedlocal water control factors, limitations in thehousehold and village-level (...) seedinfrastructure, farm location in relation tothe goods and services necessary to obtainseeds, policies and programs of the Departmentof Agriculture, and the characteristics of therice varieties themselves. The implications ofour study are that genetic erosion is notalways the result of purposeful acts by farmersnor is it necessarily gradual. Improvingon-farm seed technology will stabilize the seedproduction, distribution, and use system andthereby enhance household food security.Ultimately, rice diversity will be improvedonly if diversity is a safe and viable optionfor farmers. Therefore, public policy thatsupports farmers who maintain a diverse set ofcultivars is critical for any on-farmconservation strategy. (shrink)

In a previous issue of Tijdschrift voor Filosofie, Filip Buekens argues that evolutionary psychology (EP), or some interpretations thereof, have a corrosive impact on our ‘manifest self-image’. Buekens wants to defend and protect the global adequacy of this manifest self-image in the face of what he calls evolutionary revisionism. Although we largely agree with Buekens’ central argument, we criticize his analysis on several accounts, making some constructive proposals to strengthen his case. First, Buekens’ argument fails to target EP, because his (...) notion of the ‘constitutive conditions’ of our attitudes is too wide and too extensive. Second, his defense of the global adequacy of our attitudes does not allow for sufficient differentiation to analyze the problem of potential self-refutation with respect to EP. Third, his account of knowledge about constitutive conditions, and its impact on our self-image, is problematic. We provide a more detailed explanation for the pervasiveness of evolutionary revisionism and other misconceptions about EP. Finally, we consider in what sense EP may legitimately affect our self-image, and whether it can truly inspire corrections of our view of human nature. (shrink)

In 1990 Robert Lickliter and Thomas Berry identified the phylogeny fallacy, an empirically untenable dichotomy between proximate and evolutionary causation, which locates proximate causes in the decoding of ‘ genetic programs’, and evolutionary causes in the historical events that shaped these programs. More recently, Lickliter and Hunter Honeycutt argued that Evolutionary Psychologists commit this fallacy, and they proposed an alternative research program for evolutionary psychology. For these authors the phylogeny fallacy is the proximate/evolutionary distinction itself, (...) which they argue constitutes a misunderstanding of development, and its role in the evolutionary process. In this article I argue that the phylogeny fallacy should be relocated to an error of reasoning that this causal framework sustains: the conflation of proximate and evolutionary explanation. Having identified this empirically neutral form of the phylogeny fallacy, I identify its mirror image, the ontogeny fallacy. Through the lens of these fallacies I attempt to solve several outstanding problems in the debate that ensued from Lickliter and Honeycutt’s provocative article. (shrink)

Much good work has been done on the evolution of human morality by focusing on how “selfish genes‘ can give rise to altruistic human beings. A richer research program is needed, however, to take into account the ambivalence of naturally evolved biopsychological motivators and the historical pluralism of human morality in religious systems. Such a program is described here. A first step is to distinguish the ultimate cause of natural selection from proximate causes that are the results (...) of natural selection. Next, some proximate causes are suggested as possible conditions of biological and emotional valuing as well as of customary social morality and individual rational ethical thought. Finally, different moral perspectives of Confucianism, Hinduism, Buddhism, and Christianity are briefly presented in order to illustrate how one might inquire about the selection of a variety of biopsychological and cultural proximate causes that enable the evolution of a plurality of religious moral systems. (shrink)

Recent work on the evolution of utility has brought the blunt instrument of kin selection closer to the cluttered scalpel kit of social science. The concept of diminishing marginal utility can help streamline the latter. Reconciling ultimate causes with proximate inclinations, however, will be easier for the case of assistance from grandparents than assistance to them.

Altered attachment characteristics may disturb the HPA and can even cause changes in the hippocampus. However, it is unknown if that will influence spatial navigation performance, learning and recalling. In this study we pretend to verify if there are differences in spatial navigation learning and was associated with attachment style dimensions of anxiety and close-depend. Sixty-five female participants were recruited and were evaluated using the Adult Attachment Scale-R and tested on a virtual maze navigation task (VMT) at one moment (exploratory (...) trial + 3 trials) and 24h later (3 trials). There was a significant Moment x Trial x Close-Depend interaction for the outcome time, F (2, 126) = 3.807, p = 0.025, with post-hoc analysis indicating that the High Close-Depend group displayed significant improvements between Trial 1 and Trial 3 in the post-test assessment. Conversely, the Low Close-Depend group displayed significant improvements between Trial 1and Trial 3 but on the pre-test assessment. Furthermore, the Low Close-Depend group presented significant better performance in pre-test Trial 3 in comparison to the High Close-Depend group. Thereby, it seems that low comfort with proximity and trust in others is associated with reduced spatial navigation recall, although spatial learning performance was actually superior in these participants. It is possible that reduced exposure to social interaction and meaning relationship may be reduced in the Low Close-Depend group, leading to modifications in hippocampal function and, ultimately, reduced spatial recall. Conversely, participants with high comfort with proximity and trust in others may not display typical spatial learning in the proposed task as they are more willing to freely explore the presented environment. (shrink)

In lieu of an abstract, here is a brief excerpt of the content:164 BOOK REVIEWS Thomas Buckingham and the Contingency of Futures-The Possibility of Human Freedom: A Study and Edition of Thomas Buckingham, De contingentia futurorum et arbitrii libertate. By BARTHOLOMEW R. DE LA TORRE, O.P. University of Notre Dame, The Medieval Institute Publications in Medieval Studies, Vol. XXV. Notre Dame, Indiana: University of Notre Dame Press, 1987. Pp. xii +394. In this volume, Fr. Batholomew de la Torre offers the (...) text and an extended study of Thomas Buckingham's De contingentia futurorum et arbitrii libertate, which is the first of six questions of this scholastic's Quaestiones theologicae, more properly entitled Ostensio meriti liberae actionis. The six questions are found in their entirety in two English manuscripts, namely Oxford New College 134 and Oxford Merton 143. The New CoUege (late fourteenth century) manuscript is earlier and superior to Merton (early fifteenth century), and accordingly the editor adopts it except in cases where the Merton reading makes better sense. The author, Thomas Buckingham, born in the early days of the fourteenth century, was an Oxford scholar at Merton, chancellor of Exeter, and died, very likely a victim of the Black Death, in 1348. The work edited here probably formed a part of Buckingham's exercise prescribed by University statute for those incepting as Masters of Theology. In the editor's view, this inception took place sometime between 1341 and 1346, when Thomas departed Oxford to take up duties at Exeter. There is some reason to believe that Buckingham may have been a student of Thomas Bradwardine (p. 7); irt is certain, however, that the present work "is a sustained attack on Bradwardine's notion that all future contingents come about by antecedent necessity relative to the first cause, God" (p. 103) • Bradwardine presented his view regarding how we must reconcile God's universal and immutable causality with human freedom in his magnum opus De causa Dei contra Pelagium. De la Torre describes Bradwardine's position this way: "For Bradwardine, God is the universal cause of all things, including future contingents. Future contingents are determined by divine antecedent necessity, for God's will concerning them is omnipotent and unimpedible. Such futures nevertheless remain contingent because only man as a secondary cause-and no other secondary cause-is responsible for his free choice. The theory that, once God has determined what a future shall be, He can still determine otherwise, is patently untenable BOOK REVIEWS 165 for Bradwardine because it predicates mutability of God and nullifies all inspired prophecy, including that of Christ" (p. 101). Bradwardine's exposition here outlined corresponds in its essentials with what Thomas Aquinas had taught in the previous century. The freedom or contingency in man's will act is to he understood in terms of its relationship to secondary or proximate causes. Unlike those events which are determined to come about by virtue of necessitating proximate causes, will acts in this respect remain totally undetermined. Along with this, says Aquinas, we must allow that God, because of His eternal knowledge, knows all futures-determined, as well as contingent -as present. His knowing future contingents, however, does not take away the condition of contingency, since the latter condition rests solely on their mode of occurrence as they relate to secondary or proximate causes. Thomas Buckingham finds this explanation inadequate. In order to safeguard future contingents or human freedom. Buckingham insists there must be a sense in which the ultimate cause, God, is casually contingent with respect to the human will act. Though his exposition is different in many details, Buckingham's fundamental direction is in accord with what certain predecessors, especially Scotus and Ockham, had argued regarding the problem of divine foreknowledge and future contingents. Indeed, in view of the way Buckingham cites Scotus, the editor considers it reasonable to suppose that he regarded himself a disciple of the Subtle Doctor (p. 132). There is at least one instance where Buckingham exhibits a markedly Scotistic and anti-Ockhamist analysis. This occurs in connection with the explanation as to how in one and the same instant, because of the different moments of natural priority and posteriority, the divine will can both will and not will the same thing. The... (shrink)

Niedenthal et al's classification of smiles erroneously conflates psychological mechanisms and adaptive functions. This confusion weakens the rationale behind the types of smiles they chose to individuate, and it obfuscates the distinction between the communicative versus denotative nature of smiles and the role of perceived-gaze direction in emotion recognition.

Partner preferences are expressed by many social species, including humans. They are commonly observed as selective contacts with an individual, more time spent together, and directed courtship behavior that leads to selective copulation. This review discusses the effect of conditioning on the development of heterosexual and homosexual partner preferences in rodents. Learned preferences may develop when a conditioned stimulus (CS) is associated in contingency with an unconditioned stimulus (UCS) that functions as a reinforcer. Consequently, an individual may display preference for (...) a partner that bears a CS. Some UCS may be more or less reinforcing, depending on when they are experienced, and may be different for males and females. For example, it could be that, only during periods of early development, that stimuli associated with nurture and juvenile play become conditioned. In adulthood, other stimuli such as sexual reward, cohabitation, mild stress, or even pharmacological manipulations may function as reinforcers to condition partner preferences. Evolutionary biologists and psychologists must take into consideration the idea that an individual’s experience with reward (i.e. sexual and pharmacological) can override presumably ‘innate’ mate choices (e.g. assortativeness and orientation) or mate strategies (e.g. monogamy or polygamy) by means of Pavlovian and operant contingencies. In fact, it is likely as innate to learn about the environment in ways that maximize reward and minimize aversive outcomes, making so-called ‘proximate’ causes (e.g. pleasure) ultimately more powerful predictors of social behavior and choice than so-called ‘ultimate’ causes (e.g. genetic or reproductive fitness). Keywords: pavlovian; operant; learning; sex; copulation (Published: 15 March 2012) Citation: Socioaffective Neuroscience & Psychology 2012, 2 : 17340 - DOI: 10.3402/snp.v2i0.17340. (shrink)

These are reflections on the Arkansas creationist trial by a witness for the American Civil Liberties Union. The following points are stressed: First, religion took the lead in defending science at the trial. Second, the appearance of creation science is a function not only of Protestant fudamentalism but also of the establishment of science in our wider culture. It represents a “deviant science” in such a culture. Third, our century has manifested many such bizarre unions of ideological religion and modern (...) science. This shows that science is dependent upon its humanistic, moral, and religious matrix for its social and historical health. Fourth, part of the cause of the rise of creation science has been the power, status, and self–assurance of science that it represents “the only form of truth.” Fifth, religion in turn tends both to increase and to become fanatical in advanced and precarious cultures; religion, therefore, needs rational and moral criticism if it would help in the creation of social health. (shrink)

One of the most important issues in Islamic law is that either partially or completely, or temporary or permanently, a rule can be changed for a particular group of people or everyone. Since the concept of necessity can lead to a change of an important rule like ḥarām/prohibition, this concept should be examined meticulously both in theory and in practice. The thşs study aims to analyze how and why necessities make some ḥarāms permissible and to reveal the ultimate cause (...) for this ibāḥa (permission) from the point of view of Islamic legal theory and philosophy. To achieve this aim, first, the nature of the relationship between the prohibition and the act is examined. To understand how and why necessity gives permission to some ḥarāms, it is necessary to determine the changing and unchanging elements in the case at issue. In the case of necessity, there is no change in the nature of the act and the evil, which is considered as the reason of banning. Indeed, there is a change in the conditions surrounding the act and in the state of the obligated person. However, the main factor that enables the change in ḥarām is the change in the addressing of Shāri‘, who takes into account the changes mentioned. It is seen that the changes stemming from necessity do not lead to the same results as in all permitted ḥarāms. This observation requires a thourgh examination and elaboration on what the scholars of legal theory mean by abolition of prohibition and turning it into permissible. As a result, it has been observed that permitted ḥarāms do not equally function in terms of turning into permissible and that they do not have an equal effect on other acts in the field of mubāḥ. In that case, it is possible to evaluate the permitted ḥarām acts as al-maʻfuw ʻanh/forgiven acts. Although the act could be considered part of the category of mubāḥ, it would be more correct to place them close to the line of makrūh. From the point of view of the servant, the reason why necessity makes a ḥarām act permissible is the priority of avoiding greater evil. However, to reveal His sincere servants, Allah's command to them to avoid all ḥarāms even if they die, does not contradict with His wisdom. For this reason, the ultimate cause why Allah considers the case of necessity in passing judgment is His endless mercy. All these points helped reaching some broader conclusions about legal theory, as well as some conclusions specific to the case of necessity. It turns out that to understand how necessity makes some ḥarāms permissible, one needs to comprehend the relation between the act, the obligated person, the conditions surrounding them and the Shari‘ correctly. Understanding this relationship well is also extremely important to understand the theory of ruling and the Islamic legal theory. This research has substantiated once again that the issues addressed by the Islamic legal theory and practice are linked to the theological doctrines, more than, it is commonly perceived or considered. It turns out that it is important to know the deep or subtle differences between the terms such as ḥarām-ḥalāl, maḥẓūr-ḥarām, mubāḥ-ḥalāl, fundamental mubāḥ-ma‘fuw ‘anh, etc. as well as grasping each of the rule categories. Comprehending all these concepts constitutes an important step in acquiring a true understanding of religion and the maqāṣid al-sharī‘a. In this research, it has been proven once again that each category of the rule includes different levels of ruling in itself. The reason why the acts are categorized in different levels even if they are in the same rule category is the concepts of worldly and otherworldly; evil or good arising from the the ḥarām act. This study also confirmed that the obligated person is at the center of the law. This fact reveals how difficult it is to argue that the rules of Allah are based on divine will that is independent of and disregards the act, obligated person, and conditions. On the contrary, it is understood that the interests of the servant are considered both in the rules of necessity and in the whole of the divine legislation (and even the genesis). On the other hand, this indicates a legal system that is not inflexible and dull, but rather dynamic and parallel to life. Both the fact that the real and absolute source of the rule is Shāri‘ Almighty,and that a very large area of legislative was left to the mujtahid should be considered together. This truth has been observed once again in the matter of making prohibitions permissible on the basis of necessity, and the task of determining which evil is greater was often left to the mujtahid’s consideration. (shrink)

To understand the consequences of cleansing, Lee and Schwarz favor a grounded procedures perspective over recently developed disgust theory. We believe that this position stems from three errors: (1) interpreting cleansing effects as broader than they are; (2) not detailing the proximate mechanisms underlying disgust; and (3) not detailing adaptive function versus system byproducts when developing the grounded procedures perspective.