Elisabeth Lloyd

Two major clarifications have greatly abetted the understanding and fruitful expansion of the theory of natural selection in recent years: the acknowledgment that interactors, not replicators, constitute the causal unit of selection; and the recognition that interactors are Darwinian individuals, and that such individuals exist with potency at several levels of organization (genes, organisms, demes, and species in particular), thus engendering a rich hierarchical theory of selection in contrast with Darwin’s own emphasis on the organismic level. But a piece of the argument has been missing, and individuals at levels distinct from organisms have been denied potency (although granted existence within the undeniable logic of the theory), because they do not achieve individuality with the same devices used by organisms and therefore seem weak by comparison. We show here that different features define Darwinian individuality across scales of size and time. In particular, species-individuals may develop few emergent features as direct adaptations. The interactor approach works with emergent fitnesses, not with emergent features; and species, as a consequence of their different mechanism for achieving individuality (reproductive exclusivity among subparts, that is, among organisms), express many effects from other levels. Organisms, by contrast, suppress upwardly cascading effects, because the organismic style of individuality (by functional integration of subparts) does not permit much competition or differential reproduction of parts from within. Species do not suppress the operation of lower levels; such effects therefore become available as exaptations conferring emergent fitness—a primary source of the different strength that species achieve as effective Darwinian individuals in evolution.

This chapter explores two of the most studied and most damaging aspects of such societal influence on science: racial and gender biases. We discuss two major domains of biological and medical research involving race and gender: cognitive differences research and reproductive health science. In each case, we explore the influence of sexist values like androcentric bias—where researchers focus on men and male bodies as the alleged “norm”—and racist values like white supremacy—where researchers privilege the cultures and attributes of white people as allegedly “superior” to those of people of color. The chapter contends that we must make the culture of research more equitable to make the research itself less biased and more fair for all people.

Holobionts, consisting of a host and diverse microbial symbionts, function as distinct biological entities anatomically, metabolically, immunologically, and developmentally. Symbionts can be transmitted from parent to offspring by a variety of vertical and horizontal methods. Holobionts can be considered levels of selection in evolution because they are well-defined interactors, replicators/reproducers, and manifestors of adaptation. An initial mathematical model is presented to help understand how holobionts evolve. The model offered combines the processes of horizontal symbiont transfer, within-host symbiont proliferation, vertical symbiont transmission, and holobiont selection. The model offers equations for the population dynamics and evolution of holobionts whose hologenomes differ in gene copy number, not in allelic or loci identity. The model may readily be extended to include variation among holobionts in the gene identities of both symbionts and host.

We critically examine Denis Walsh’s latest attack on the causalist view of fitness. Relying on Judea Pearl’s Sure-Thing Principle and geneticist John Gillespie’s model for fitness, Walsh has argued that the causal interpretation of fitness results in a reductio. We show that his conclusion only follows from misuse of the models, that is, (1) the disregard of the real biological bearing of the population-size parameter in Gillespie’s model and (2) the confusion of the distinction between ordinary probability and Pearl’s causal probability. Properly understood, the models used by Walsh do not threaten the causalist view of fitness.

climate scientists have been engaged in a decades-long debate over the standing of satellite measurements of the temperature trends of the atmosphere above the surface of the earth. This is especially significant because skeptics of global warming and the greenhouse effect have utilized this debate to spread doubt about global climate models used to predict future states of climate. I use this case from an under-studied science to illustrate two distinct philosophical approaches to the relation among data, scientists, measurement, models, and theory. I argue that distinguishing between 'direct' empiricist and 'complex' empiricist approaches helps us understand and analyze this important scientific episode. I also introduce a complex empiricist account of testing and evaluation, and contrast it with the basic Hypothetico-Deductive approach to the climate models used by the direct empiricists. This more developed complex empiricist apporach will serve philosophy of science well, as computational models become more wide-spread in the sciences.

Science strives for coherence. For example, the findings from climate science form a highly coherent body of knowledge that is supported by many independent lines of evidence: greenhouse gas emissions from human economic activities are causing the global climate to warm and unless GHG emissions are drastically reduced in the near future, the risks from climate change will continue to grow and major adverse consequences will become unavoidable. People who oppose this scientific body of knowledge because the implications of cutting GHG emissions—such as regulation or increased taxation—threaten their worldview or livelihood cannot provide an alternative view that is coherent by the standards of conventional scientific thinking. Instead, we suggest that people who reject the fact that the Earth’s climate is changing due to greenhouse gas emissions oppose whatever inconvenient finding they are confronting in piece-meal fashion, rather than systematically, and without considering the implications of this rejection to the rest of the relevant scientific theory and findings. Hence, claims that the globe “is cooling” can coexist with claims that the “observed warming is natural” and that “the human influence does not matter because warming is good for us.” Coherence between these mutually contradictory opinions can only be achieved at a highly abstract level, namely that “something must be wrong” with the scientific evidence in order to justify a political position against climate change mitigation. This high-level coherence accompanied by contradictory subordinate propositions is a known attribute of conspiracist ideation, and conspiracism may be implicated when people reject well-established scientific propositions.

The theory of evolution by natural selection is, perhaps, the crowning intellectual achievement of the biological sciences. There is, however, considerable debate about which entity or entities are selected and what it is that fits them for that role. This article aims to clarify what is at issue in these debates by identifying four distinct, though often confused, concerns and then identifying how the debates on what constitute the units of selection depend to a significant degree on which of these four questions a thinker regards as central

I admire Wilson & Sober's (W & S's) aim, to alert social scientists that group selection has risen from the ashqs, and to explicate its relevance to the behavioral sciences. Group selection has beenwidely misunderstood; furthermore, both authors have been instrumental in illuminating conceptual problems surrounding higher-level selection. Still, I find that this target article muddies the waters, primarily through its shifting and confused definition of a "vehicle" of selection. The fundamental problem is an ambiguity in the definition of "adaptation." On the one hand, any evolutionary change that results from a selection process could be called an adaptation, by definition; I call this the "weak" view of adaptation. A "strong" view of adaptation, on the other hand, includes some notion of design - the evolution of a specific complex trait understood, in an engineering sense, to provide a mechanism favoring its owner's success in contributing to the evolutionary lineage.

Some methodological adaptationists hijacked the term “exaptation,” and took an occasion of Stephen Jay Gould’s misspeaking as confirmation that it possessed an evolutionarily “designed” function and was a version of an adaptation, something it was decidedly not. Others provided a standard of evidence for exaptation that was inappropriate, and based on an adaptationist worldview. This article is intended to serve as both an analysis of and correction to those situations. Gould and Elisabeth Vrba’s terms, “exaptation” and “aptation,” as originally introduced, are very useful, unlike the faded adaptationist echo of “exaptation” devised by the methodological adaptationists, which has made the term incoherent. We will discuss how exaptation relates to function, to aptation, and to adaptation, both primary and secondary. These ideas have been rendered practically useless through their mistaken definitions and misapplications by evolutionary psychologists.

1. Introduction; Elisabeth A. Lloyd and Eric Winsberg.- Section 1: Confirmation and Evidence.- 2. The Scientific Consensus on Climate Change: How Do We Know We’re Not Wrong?; Naomi Oreskes.- 3. Satellite Data and Climate Models Redux.- 3a. Introduction to Chapter 3: Satellite Data and Climate Models; Elisabeth A. Lloyd.- Ch. 3b Fact Sheet to "Consistency of Modelled and Observed Temperature Trends in the Tropical Troposphere"; Benjamin D. Santer et al..- Ch. 3c Reprint of "Consistency of Modelled and Observed Temperature Trends in the Tropical Troposphere"; Benjamin D. Santer et al..- 4. The Role of ’Complex’ Empiricism in the Debates about Satellite Data and Climate Models; Elisabeth A. Lloyd.- 5. Reconciling Climate Model/Data Discrepancies: The Case of the Trees That Didn’t Bark; Michael Mann.- 6. Downscaling of Climate Information; Linda O. Mearns et al..- Section 2: Uncertainties and Robustness.- 7. The Significance of Robust Model Projections; Wendy S. Parker.- 8. Building Trust, Removing Doubt? Robustness Analysis and Climate Modeling; Jay Odenbaugh.- Section 3: Climate Models as Guides to Policy.- 9. Climate Model Confirmation: From Philosophy to Predicting Climate in the Real World; Reto Knutti.- 10. Uncertainty in Climate Science and Climate Policy; Jonathan Rougier and Michel Crucifix.- 11. Communicating Uncertainty to Policy Makers: The Ineliminable Role of Values; Eric Winsberg.- 12. Modeling Climate Policies: A Critical Look at Integrated Assessment Models; Mathias Frisch.- 13. Modelling Mitigation and Adaptation Policies to Predict their Effectiveness: The Limits of Randomized Controlled Trials; Alexandre Marcellesi and Nancy D. Cartwright.

A specific form of research question, for instance, “What is the probability of a certain class of weather events, given global climate change, relative to a world without?” could be answered with the use of FAR or RR (Fraction of Attributable Risk or Risk Ratio) as the most common approaches to discover and ascribe extreme weather events. Kevin Trenberth et al. (2015) and Theodore Shepherd (2016) have expressed doubts in their latest works whether it is the most appropriate explanatory tool or the way of public outreach concerning climate events and extremes. As an alternative, these researchers focus on complementary questions, for example, “How much did climate change affect the severity of a given storm?” advocating a “storyline” approach. New methods and new research questions are neither foreign, nor controversial from the standpoint of history and philosophy of science, especially those, related to public interest. Nevertheless, the new proposal has got a tepid reception from the majority of professionals of the given field. They argued that this approach can cause weakening of standards and neglecting of scientific method. The following paper attempts to find the roots of the supposed controversy. We claim inefficiency of uncompromising approach to D&A in absolute sense and assert that errors of a particular type may have a different level of concern in society, depending on the variety of contexts. Therefore, context defines the risk of over-estimation vs. under-estimation of harm.

In this talk I do three things. First, I review what I take to be fruitful applications of the semantic view of theory structure to evolutionary theory. Second, I list and correct three common misunderstandings about the semantic view. Third, I evaluate the weaknesses and strengths of Horan's paper in this symposium. Specifically, I argue that the criticisms leveled against the semantic view by Horan are inappropriate because they incorporate some basic misconceptions about the semantic view itself.

We address the controversy in the literature concerning the definition of holobionts and the apparent constraints on their evolution using concepts from community population genetics. The genetics of holobionts, consisting of a host and diverse microbial symbionts, has been neglected in many discussions of the topic, and, where it has been discussed, a gene-centric, species-centric view, based in genomic conflict, has been predominant. Because coevolution takes place between traits or genes in two or more species and not, strictly speaking, between species, it may affect some traits but not others in either host or symbiont. Moreover, when interacting species pairs are embedded in a larger community, indirect ecological effects can alter the expected pairwise dynamics. Mode of symbiont transmission and the degree of host inbreeding both affect the extent of microbial mixing across host lineages and thereby the degree to which selection on one trait of either partner affects other aspects of a holobiont phenotype. We discuss several potential defining criteria for holobionts using community genetics and population genetics models, suggesting their application and limitations. Using community genetics models, we show how conflict between genomes can be self-limiting, while cooperation and mutualism tend to be self-accelerating. It is likely that this bias in the evolutionary dynamics of interaction between hosts and symbionts is an important feature of holobionts. This bias in the evolutionary dynamic could contribute to explaining the absence of cheaters from natural mutualisms, although cheaters are predicted by gene-centered conflict theory to cause the evolutionary instability of mutualisms. Additionally, it may help explain the more frequent origin of mutualisms from parasitic than from free-living systems, an evolutionary trajectory opposite to that predicted by genome conflict theory.

I address the controversy in evolutionary biology concerning which levels of biological entity (units) can and do undergo natural selection. I refine a definition of the unit of selection, first presented by William Wimsatt, that is grounded in the structure of natural selection models. I examine Elliott Sober's objection to this structural definition, the "homogeneous populations" problem; I find that neither the proposed definition nor Sober's own causal account can solve the problem. Sober, in his solution using his causal view, imports precisely the information needed to make the structural definition effective. Finally, I indicate how the proposed definition can clarify which sorts of evidence could be brought to bear on the controversial case of the Myxoma virus.

Elisabeth Lloyd is an American philosopher of science whose work is centered in the field of philosophy of biology. The material in this archive documents her work in philosophy of biology. The materials extend over the whole of her career and include manuscript materials, working notes on articles and books in progress, professional correspondence, teaching materials, documents relating to work with professional organizations, talks given to professional audiences, as well as annotated books, manuscripts and preprints. Elisabeth Lloyd's publications include both books and professional articles.

Feminist philosophers have discussed the prospects for assessing values empirically, particularly given the ongoing threat of sexism and other oppressive values influencing science and society. Some advocates of such tests now champion a “values as evidence” approach, and they criticize Helen Longino’s contextual empiricism for not holding values to the same level of empirical scrutiny as other claims. In this paper, we defend contextual empiricism by arguing that many of these criticisms are based on mischaracterizations of Longino’s position, overstatements of certain claims, and false dichotomies. Her contextual empiricism not only allows for the empirical support and disconfirmation of values, but Longino explicitly discusses when values can be empirically adjudicated and emphasizes the crucial role of the community for standards of evidence. We support contextual empiricism and elaborate a less direct account of “values as heuristics” by reviewing Longino’s theory of evidence and then using a case study from Elisabeth Lloyd on the biology of female orgasm, demonstrating the disconfirmation of androcentric values in evolutionary science. Within Longino’s and Lloyd’s contextual empiricism, values do not get treated as empirical evidence to be directly assessed by individuals, but rather values are heuristic tools to build models whose use can be validated or invalidated by communities based on their empirical fruitfulness in the logic and pragmatics of research questions in specific historical and cultural contexts.

Most models of generational succession in sexually reproducing populations necessarily move back and forth between genic and genotypic spaces. We show that transitions between and within these spaces are usually hidden by unstated assumptions about processes in these spaces. We also examine a widely endorsed claim regarding the mathematical equivalence of kin-, group-, individual-, and allelic-selection models made by Lee Dugatkin and Kern Reeve. We show that the claimed mathematical equivalence of the models does not hold.

This book brings together important essays by one of the leading philosophers of science at work today. Elisabeth A. Lloyd examines several of the central topics in philosophy of biology, including the structure of evolutionary theory, units of selection, and evolutionary psychology, as well as the Science Wars, feminism and science, and sexuality and objectivity. Lloyd challenges the current evolutionary accounts of the female orgasm and analyses them for bias. She also offers an innovative analysis of the concept of objectivity. Lloyd analyses the structure of evolutionary theory and unlocks the puzzle of the units of selection debates into four distinct aspects, illuminating several mysteries in the biology literature. Central to all essays in this book is the author's abiding concern for evidence and empirical data.