natural order research paper

2005 Articles

The “Natural Order” of Morpheme Acquisition: A Historical Survey and Discussion of Three Putative Determinants

Kwon, Eun-Young

The paper discusses the history and implications of the so-called “morpheme” studies. A brief overview of L1 research in this regard is followed by a more detailed discussion of L2 research. Thereafter, the paper explores the impact of three putative determinants of acquisition order: semantic complexity, input frequency, and native language transfer. The possible role of these determinants in accounting for perceived differences in L1 and L2 acquisition orders is also discussed, along with their implications for various theoretical perspectives on language acquisition. The paper concludes with both practical applications and criticism of existing natural (morpheme) order studies as well as suggestions for future work in this field, such as investigating target languages other than English and developing a multi-determinant approach to acquisition order.

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First things first: The pragmatics of “natural order”

Laurence Horn is Professor Emeritus of Linguistics and Philosophy at Yale University. He is the author of A Natural History of Negation (Chicago, 1989; CSLI, 2001) and over 100 papers and handbook entries on negation, polarity, implicature, presupposition, pragmatic theory, word meaning, grammatical variation, and lying. His PhD dissertation (UCLA, 1972) introduced scalar implicature. His six (co-)edited volumes include The Handbook of Pragmatics (Blackwell, 2004; co-edited with Gregory Ward) and Pragmatics, Truth and Underspecification (Brill, 2018; co-edited with Ken Turner). He is an elected fellow of the Linguistic Society of America and edited the Outstanding Dissertations in Linguistics series (Garland/Routledge).

Classical rhetoricians dating back to Aristotle sought to define the principles of natural order that determine priority in sequences, especially in linguistic representations. Among the principles with the widest predictive power for the ancients and their modern heirs are those stating that A can be prior to B “with respect to temporal order”, that A can be prior to B with respect to what is “known or less informative” than what comes later, and that A can be prior to B with respect to what is “better” or “more worthy”. But when and how do these ordering principles influence the form of linguistic sequences, and how are conflicts between the principles resolved? What determines the priority between the principles of priority? What makes “natural order” natural? Drawing on over two millennia of scholarship, we explore the pragmatic motivation for the primary ordering principles, and in particular for those affecting the order of logically symmetric but rhetorically asymmetric conjunctions.

About the author

Acknowledgements.

“First Things First” was presented at Pragmasophia 2 (Lisbon, 2018). Earlier versions were given at CIL 19 (Geneva, 2013), at the CUNY Pragmatics Workshop: Relevance, Game Theory, and Communication (2014), at AMPRA [American Pragmatics Association] 2 (UCLA, 2014), and the University of Cologne (2016). I thank (better late than never!) the audiences at those presentations (and in particular Istvan Kecskes, Klaus von Heusinger, and Petra Schumacher) as well as Barbara Abbott, Mira Ariel, Betty Birner, Bob Frank, Michael Israel, Paul Kay, Gregory Ward, and Ben Zimmer for judgments, discussion, and encouragement. Empirical work on reverse substitute was supported in part by NSF grant BCS-1423872: The Microsyntax of Pronouns in North American English; thanks to Jim Wood and Raffaella Zanuttini, and other collaborators on the grant. I am indebted to the American Dialect Society e-mail list, the Yale Grammatical Diversity group, and the participants in the University of Geneva winter school (Dec. 2015) in Macolin, Switzerland for data and feedback and to Arnold Zwicky and David Denison for helpful guidance. Thanks to Huber (1974 ), a work I have not been able to consult, for providing the Section § 4 header. Last but not least (#not least but last), everyone but me is absolved of any errors.

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Neoliberal Penality: The Birth of Natural Order, the Illusion of Free Markets

U of Chicago Law & Economics, Olin Working Paper No. 433

U of Chicago, Public Law Working Paper No. 238

69 Pages Posted: 6 Oct 2008 Last revised: 13 Jan 2009

Bernard E. Harcourt

Columbia University

Date Written: October 2, 2008

This Article represents the culmination of over two-years of historical research, but it arrives at an odd moment, right in the middle of one of the largest financial crises in Western capitalism. In one sense, it is bad timing because the central premise of the Article is that most people today believe that the market is the most efficient mechanism to allocate resources. The federal bailouts of 2008 challenge this central premise and are forcing the American people to reexamine the need for the regulation of the free market. In another sense, the timing is, sadly, perfect. Perfect because the purpose of this Article is to question the meaning of the phrase the need for the regulation of the free market and to suggest that it is precisely the belief in the duality of those two terms - regulation and free market - that is one of the greatest problems we face today. The terms, as well as their companion expressions, market efficiency, natural order, self-adjusting markets, etc., are misleading categories that fail to capture the individual distinctiveness of different forms of market organization. These categories are responsible, first, for facilitating our growing penal sphere, and, second, for naturalizing and thereby masking the redistributive consequences associated with different methods of organizing markets. This Article asks the question, what work do these categories of natural order and market efficiency do for us? The story begins very far in time and place, in the Parisian markets of the eighteenth century, with the establishment of the lieutenant generale de police du Chatelet de Paris and the police of bakers, grain merchants, and markets.

Keywords: Free market, regulation, market efficiency, efficiency, natural order, punishment, neoliberalism, neoliberal, mass incarceration, policing, Chicago Board of Trade, grain markets, regulated markets, free trade, regulated trade, Parisian bread markets, self-regulation, self-adjusting markets, legal de

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3 Laws of God or Laws of Nature? Natural Order in the Early Modern Period

• Published: January 2019
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The appeal to laws of nature as an explanatory principle is often regarded as fundamental to naturalism. Yet when the idea that there were immutable, mathematical laws of nature first rose to prominence in the seventeenth century it was deeply connected to a theological understanding of natural order. Descartes thus imagined laws of nature to be divine commands, and attributed their immutability to the immutability of their divine source. For Descartes, Boyle, and Newton, the invariable uniformity of nature was understood as a consequence not of God’s withdrawal from the world, but of his direct and incessant engagement with it. It followed that the world was to be investigated empirically, because this was the only way in which the otherwise inscrutable will of God could be discerned. Over the course of the following centuries, however, laws came to be reimagined as simply observational generalizations, or brute features of the natural world.

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• Corpus ID: 142087074

Natural order : historical studies of scientific culture

• B. Barnes , S. Shapin
• Published 1979
• History, Sociology

174 Citations

Cultural anthropology and the paradigm-concept: a brief history of their recent convergence, on the relationship of the criticism of ethnographic writing and the cultural studies of science, cultural history of science, cultural arts research and development, three genres of sociology of knowledge and their marxist origins, facing gaia: eight lectures on the new climatic regime, the scope of hermeneutics in natural science, the politics of truth: a social interpretation of scientific knowledge, with an application to the case of sociobiology, sociology and the mirror of nature: robert brandom and the strong programme, related papers.

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Series on the Foundations of Natural Science and Technology: Volume 12

• The Origin of Natural Order
• By (author): 
• Qinyi Zhao ( Medical Institute of China Rehabilitation Research Centre, China )
• Add to favorites
• Download Citations
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• Description
• Supplementary

All sorts of biological activities are processed thermodynamically, and at the utmost fundamental level, the laws of biology must be thermodynamics. However, the current laws of thermodynamics are unable to give reasonable explanation of biological processes. In order to do so, irreversible thermodynamics has been theorized to describe the basic mechanism for the origin of natural order or the development of things (related to developmental biology). The scientific definition of the system theory concept has been obtained and the properties of a biological system can be analyzed by applying principles of it. Irreversible thermodynamics and system theory act as the theoretical foundation for theoretical biology. By applying principles of irreversible thermodynamics and system theory, the axiomatic theory of biology has been developed.

Chinese poster of the book (215 KB)

Sample Chapter(s) Introduction (85 KB) Chapter 1: The Essence of Irreversible Thermodynamics (180 KB)

• Introduction
• The Essence of Irreversible Thermodynamics
• The Developing System and Hierarchical Structure of Things
• Partition Function of a Thermodynamic System
• Dissipative Structure and Thermodynamic Structure Theory
• The Scientific Definition of the System
• Logic Discontinuity and Continuity Between Different Levels of Hierarchical Structure
• System Logic
• Isolating Logic Relationship Between Complex Systems
• Irreversible Thermodynamic Structure Theory for Protein Folding
• Can We Go from Protein Sequence to Protein Structure?
• Protein Thermodynamic Structure Theory
• General Properties of a Protein
• Protein Conformation
• Dynamic and Thermodynamic Nature of Protein Conformational Change
• Protein Ligand Interaction
• The Receptor Activation and Fashions of Protein Regulation
• The Basic Thermodynamic Principles of Enzyme Catalyzed Reaction
• Protein Conformational Change and Enzyme Activity
• Allodynamic Regulation Model for Enzyme Regulation — A Thermodynamic View
• The Protein Evolution
• A Thermodynamic Model for Bio-signals
• Protein Flexibility as a Biosignal
• Signal Conduction and Neural Conduction
• Signaling Network Theory and System Biology
• Biological Development
• Hybridization and Hybrid Vigor
• Biological Evolution
• Scientific Explanation of Traditional Chinese Medical Theory

FRONT MATTER

• Pages: i–xv

https://doi.org/10.1142/9789813209275_fmatter

Chapter 1: The Essence of Irreversible Thermodynamics

• Pages: 3–16

https://doi.org/10.1142/9789813209275_0001

Summary: The basic concept of irreversible thermodynamics is discussed in this chapter. We show that in all movements (processes) of irreversible thermodynamics, there is a giant space in which all changes of thermodynamics obey the principle of equilibrium thermodynamics, but their features cannot be fully explained by it. The new order can be generated in the processes of irreversible thermodynamics. Thus, the origin of natural order can be fully explained by its principle.

Chapter 2: The Origin of Natural Order

• Pages: 17–26

https://doi.org/10.1142/9789813209275_0002

Summary: In irreversible thermodynamics, the generation of a sub-system within a thermodynamics system represents the fundamental mechanism for the origin of natural order. According to this theory, the origin of natural order is the most common phenomenon of nature. This new understanding makes us analyze biology theoretically.

Chapter 3: The Developing System and Hierarchical Structure of Things

• Pages: 27–31

https://doi.org/10.1142/9789813209275_0003

Summary: The production of irreversible process is a hierarchical structure. If many types of hierarchical structure can be generated within a system, it is developing system. The nature and living body are developing systems.

Chapter 4: Partition Function of a Thermodynamic System

• Pages: 33–42

https://doi.org/10.1142/9789813209275_0004

Summary: The partition function is a fundamental concept of equilibrium thermodynamics. In the views of traditional thermodynamics, it encodes the particle distribution at different energy levels. However, the behaviors of the complex system of thermodynamics cannot be described by the partition function of a particle. By modifying the partition function of a particle, the partition function of a thermodynamic system has been obtained. It could be utilized to analyze behaviors of a complex thermodynamic system, such as protein conformational state, DNA conformational state. The typical fashions of distributions curves of a thermal system are of great importance and we can judge the properties of a system on the basis of it.

Chapter 5: Dissipative Structure and Thermodynamic Structure Theory

• Pages: 43–50

https://doi.org/10.1142/9789813209275_0005

Summary: In view of thermodynamics, all actual changes of nature are irreversible, and our understanding of nature, as well science, is established on the basis of principles of irreversible thermodynamics. Currently there are two different approaches in the field. First is the theory of dissipative structure which represents a dynamic but steady state of a system. Second is thermodynamic structure which represents a static state of a thermal system. The key task of this chapter is to ascertain the validation domains and different usefulness of both theories. The advantages of both theories in diversified area of science are also discussed.

Chapter 6: The Scientific Definition of the System

• Pages: 53–60

https://doi.org/10.1142/9789813209275_0006

Summary: The system is a basic concept of science. However, until now, there has been scientific definition about it. In this chapter, we discuss its scientific definition on the basis of thermodynamics. According to this definition, a system represents one specific ensemble of matters with its specific structure: logic cycle.

Chapter 7: Logic Discontinuity and Continuity Between Different Levels of Hierarchical Structure

• Pages: 61–67

https://doi.org/10.1142/9789813209275_0007

Summary: We show that the fundamental properties of matter cannot logically determine the properties of advanced matter. There is logical discontinuity between different levels of hierarchical structure, which reveals the theoretical limit of scientific model or paradigms. It also shows that we cannot analyze advanced matter by fundamental properties alone.

Chapter 8: System Logic

• Pages: 69–79

https://doi.org/10.1142/9789813209275_0008

Summary: All things in nature are systems, but the principles of system have not been formulated mathematically. Here, we formulate a standard paradigm for systems. Basic concept: 1) a system is complex system with its particular structure and it cannot be described by element logic (or axiomatic theory, the traditional logic of mathematics). 2) The relationship of systems can be only described by system logic. The relationship between system logic and element logic are discussed. It provides a new way for our understanding of complexity of things.

Chapter 9: Isolating Logic Relation Between Complex Systems

• Pages: 81–85

https://doi.org/10.1142/9789813209275_0009

Summary: When complex is merged, it is impossible to find absolute model which can account for all matters. In these cases, it is valuable to find conditional models and the logical relationship between different matter could be constructed within these models. These isolated models have great impact on our understanding of nature.

Chapter 10: Irreversible Thermodynamic Structure Theory for Protein Folding

• Pages: 89–108

https://doi.org/10.1142/9789813209275_0010

Summary: When polypeptide is synthesized in vivo , the nascent peptide begins to fold itself and finally a folded protein with full biological function is produced. Theoretically speaking, the protein folding is irreversible thermodynamic process and we discuss its principles here. Key thoughts: 1) nature of protein folding can be described by laws of irreversible thermodynamics; 2) a folded protein may be at metastable state in view of thermodynamics; 3) the product of protein folding is protein thermodynamic structure; 4) the Gibbs free energy is not the controlling factor for protein folding.

Chapter 11: Can We Go from Protein Sequence to Protein Structure?

• Pages: 109–114

https://doi.org/10.1142/9789813209275_0011

Summary: A protein has its unique three-dimensional structure and sequence. In protein science, a well-accepted view is that information of protein three dimensional structure and protein folding has been prescribed within sequence of a protein. Thus, some scientists believe that we can find a program, on the basis of which the protein three dimensional structure could be deduced from protein sequence. But the solution for it has not been obtained now. Our conclusion is that we cannot predict all protein three-dimensional structures from protein sequences using one program, nor can we predict protein function from protein three dimensional structure with one program. This represents the logic limit of protein science.

Chapter 12: Protein Thermodynamic Structure Theory

• Pages: 115–126

https://doi.org/10.1142/9789813209275_0012

Summary: A protein is not a uniform system of thermodynamics, but is composed of many sub-systems of thermodynamics, called potherse. This sub-system can be structured differently, thus comes the concept of protein thermodynamic structure. In the following chapters of this book, we will show that properties of a protein can be analyzed by applying principle of protein thermodynamic structure theory. Then this chapter will discuss basic principles of protein thermodynamic structure theory.

Chapter 13: General Properties of a Protein

• Pages: 127–134

https://doi.org/10.1142/9789813209275_0013

Summary: The structural features and the biophysical natures of a protein are discussed on the basis of protein thermodynamic structure theory. We have demonstrated that much current knowledge about protein conformation and conformational change can be explained by applying the principles of protein thermodynamic structure theory.

Chapter 14: Protein Conformation

• Pages: 135–147

https://doi.org/10.1142/9789813209275_0014

Summary: In the field of biochemistry and molecular biology, much knowledge of protein science has been explained on the basis of the concept of protein conformation. Here, we will study concept of protein conformation on the basis of protein thermodynamic structure theory. The analysis shows that a protein can show numerous types of protein conformations under different conditions.

Chapter 15: Dynamic and Thermodynamic Nature of Protein Conformational Change

• Pages: 149–159

https://doi.org/10.1142/9789813209275_0015

Summary: Each protein is a complex system of thermodynamics and its behaviours differ from that of common materials in principle. If we use our knowledge of conventional physics when studying protein science, there may be mistakes and controversies. In fact, some thermodynamic natures of protein conformational change have been classified into the dynamic nature of it. Our analysis shows that the dispute about the role of protein dynamics in enzyme activity and protein regulation results from the wrong understanding of dynamic and thermodynamic nature of protein conformational change. There is a direct relation between protein thermodynamics and enzyme activity (a type of protein functioning); the protein dynamics takes its role in enzyme activity by influencing thermodynamic properties of protein conformational states.

Chapter 16: Protein Ligand Interaction

• Pages: 161–171

https://doi.org/10.1142/9789813209275_0016

Summary: The protein ligand interaction is an important process in biology and is connected to the biological regulation of protein and enzyme. The binding between enzyme and substrate, protein-protein interaction, ligand-receptor are examples of this process. Here we analyze the basic mechanism for it on the basis of the protein conformation concept.

Chapter 17: The Receptor Activation and Fashions of Protein Regulation

• Pages: 173–188

https://doi.org/10.1142/9789813209275_0017

Summary: The receptor activation is an important process in biology. The signaling activity of a receptor is initiated at this step. Many types of biological information or signals are integrated in this process, so it has attracted much attention from scientific community in the past. Many valuable opinions about it have been proposed as well. The receptor activation is a good example of protein thermodynamic reorganization and protein regulation. In this chapter, we will show that the protein thermodynamic structure theory can give satisfactory explanations for much of the knowledge we have obtained in this field. In principle, enzyme regulation is same as the receptor activation and we will discuss some general trend of the receptor and enzyme activation.

Chapter 18: The Basic Thermodynamic Principles of Enzyme Catalyzed Reaction

• Pages: 189–198

https://doi.org/10.1142/9789813209275_0018

Summary: For simple chemical reactions, the catalyst mechanism is rather simple. However, it is more complex for an enzyme catalyzed reaction, where many chemical mechanisms are utilized at same time, which greatly reduce the activation energy of chemical reaction. This chapter will discuss the chemical mechanisms of enzymatic reaction.

Chapter 19: Protein Conformational Change and Enzyme Activity

• Pages: 199–209

https://doi.org/10.1142/9789813209275_0019

Summary: The execution of an enzyme activity requires many steps of protein conformational change by the way of substrate binding conformation, transitional state conformation and product binding conformation. If the pathway of this protein conformational change was modified, thermodynamic and dynamic parameters of enzymatic kinetics, such as k m , k cat , activation energy, as well as the affinity between enzyme and substrate will be altered. Thus, we can judge protein conformational change related to an enzyme activity by a change of parameter of enzymatic reaction. Here the basic principles in our understanding of the behaviors of enzyme will be discussed.

Chapter 20: Allodynamic Regulation Model for Enzyme Regulation — A Thermodynamic View

• Pages: 211–222

https://doi.org/10.1142/9789813209275_0020

Summary: The allosteric regulation model is a tradition model of enzyme regulation. It states that a change of protein conformation at a site can be transmitted into remote areas of the protein because the conformation of a protein can change cooperatively, which is the nature of protein three-dimensional structure. On the basis of the protein thermodynamic structure theory, an allodynamic mechanism for enzyme regulation has been proposed and it is a thermodynamic model for enzyme regulation. It states that a change of protein thermodynamic state at one site can be transmitted into remote sites of the protein via protein thermodynamic structure reorganization. A comprehensive understanding of enzyme regulation can be obtained on the basis of this model.

Chapter 21: The Protein Evolution

• Pages: 223–232

https://doi.org/10.1142/9789813209275_0021

Summary: A protein can change its properties and transform itself into another species of protein during biological evolution. This chapter discusses the basic and general principle of protein evolution.

Chapter 22: A Thermodynamic Model for Bio-signals

• Pages: 233–245

https://doi.org/10.1142/9789813209275_0022

Summary: The scientific definition of molecular bio-signals has been made on the basis of protein thermodynamics in this chapter and properties of protein bio-signals will be discussed as well. Within this model, any type of bio-signal corresponds to one potherse or a thermal system within a signaling protein. The signaling activity of a protein represents its physiological activities at a fundamental level. Any physiological activity at different levels of biology is composed of many types signaling activities of proteins.

Chapter 23: Protein Flexibility as a Biosignal

• Pages: 247–257

https://doi.org/10.1142/9789813209275_0023

Summary: Summary: Protein dynamics or flexibility plays an important role in all types of signaling activities of proteins for it alters protein conformational stability and the rate of protein conformational change. In this chapter, we will discuss some signaling activities or biological functions of protein dynamics. The experimental results have indicated that a change of protein flexibility induced by volatile anesthetics is the basis of the working mechanism for anesthesia induction.

Chapter 24: Signal Conduction and Neural Conduction

• Pages: 259–276

https://doi.org/10.1142/9789813209275_0024

Summary: The unified model of signal conduction and the protein model of neural conduction are discussed in this chapter. We will show that neural conduction is not the propagation of electric signals, but the propagation of protein conformational signals. Thus, a systematic and unified knowledge of molecular biology, neuroscience, and physiology, has been successfully developed in this chapter.

Chapter 25: Signaling Network Theory and System Biology

• Pages: 277–299

https://doi.org/10.1142/9789813209275_0025

Summary: This chapter discusses basic and thermodynamic principle of the biosignal network theory or system biology. In view of the signaling theory, a physiological system that represents a specific signaling network which is thermodynamically stable and shows independence to some degree. The properties of signaling network could be quantitatively described by the law of partition function of a thermal system. Some biological phenomena, such as plant vernalization, canceration, sexualization, and oscillation, are also highlighted in this chapter. (This chapter is a copy of the paper: A Thermodynamic Model of Protein Signaling Network. Reviews in Theoretical Science 4, pp. 1–9, 2016, with minor modification).

Chapter 26: Biological Development

• Pages: 301–313

https://doi.org/10.1142/9789813209275_0026

Summary: The general principles of biological development are discussed on the basis of the irreversible thermodynamic and signaling network theory. It is shown that the bio-informational flow from a gene to its biological function is a process of informational integration, rather than the translation process. The integration process cannot be described by a unified program (or model), thus the system relation between genes and biological function has been proposed.

Chapter 27: Hybridization and Hybrid Vigor

• Pages: 315–328

https://doi.org/10.1142/9789813209275_0027

Summary: Hybridization is a common process in biological reproduction. The change in the genetic background of an organism has a great impact on all phenotypes and biological functions of the organism. Among all the alterations, which are induced by a change in genetics, hybrid vigor is a well-established phenomenon. The working mechanism of hybridization is discussed on the basis of biological development and the signal network theory. It shows that many mechanisms have made a contribution to the production of heterosis and the main mechanism is the loose control of a physiological activities of organism.

Chapter 28: Biological Evolution

• Pages: 329–341

https://doi.org/10.1142/9789813209275_0028

Summary: From the perspective of irreversible thermodynamics, biological evolution is a specific example for the origin of natural order and is an irreversible process. The principle of biological evolution is discussed in this chapter on the basis of irreversible thermodynamics. The essential idea is that any biological species represents a specific signaling network and the behavior of biological evolution can be described and analyzed by applying the principle of system biology.

Chapter 29: Scientific Explanation of Traditional Chinese Medical Theory

• Pages: 343–359

https://doi.org/10.1142/9789813209275_0029

Summary: Traditional Chinese medical (TCM) theory developed in the ancient times based on our daily experiences and medical practice. It has incorporated these experiences into a unified philosophical theory of biology. In contrast, modern biology developed on the basis of the scientific methods, which, although extremely valuable, are unable to explain our daily experience. A scientific explanation of TCM will be valuable for expanding the scope of medical science and biology. Reconciliation between the scientific approach and TCM is therefore a key task for theoretical biology. Here, we will show that TCM can be explained by applYing principles of biological theory or system biology. The scientific account of TCM provides a comprehensive understanding of biology and medical science.

BACK MATTER

• Pages: 361–364

https://doi.org/10.1142/9789813209275_bmatter

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The Natural Order-Generic Collapse for ω -Representable Databases over the Rational and the Real Ordered Group

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• Nicole Schweikardt 5  

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We consider order-generic queries, i.e., queries which commute with every order-preserving automorphism of a structure’s universe. It is well-known that first-order logic has the natural order-generic collapse over the rational and the real ordered group for the class of dense order constraint databases (also known as finitely representable databases ). I.e., on this class of databases over 〈ℚ,〉 or 〈ℝ, 〉, addition does not add to the expressive power of first-order logic for defining order-generic queries. In the present paper we develop a natural generalization of the notion of finitely representable databases, where an arbitrary (i.e. possibly infinite) number of regions is allowed. We call these databases ω-representable , and we prove the natural order-generic collapse over the rational and the real ordered group for this larger class of databases.

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On the Expressive Power of Logics on Constraint Databases with Complex Objects

A counterexample to the reconstruction of ω-categorical structures from their endomorphism monoid.

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Nicole Schweikardt

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Schweikardt, N. (2001). The Natural Order-Generic Collapse for ω -Representable Databases over the Rational and the Real Ordered Group. In: Fribourg, L. (eds) Computer Science Logic. CSL 2001. Lecture Notes in Computer Science, vol 2142. Springer, Berlin, Heidelberg. https://doi.org/10.1007/3-540-44802-0_10

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Natural selection according to Darwin: cause or effect?

Ben bradley.

School of Psychology, Charles Sturt University, 164 George Street, Bathurst, NSW 2795 Australia

In the 1940s, the ‘modern synthesis’ (MS) of Darwinism and genetics cast genetic mutation and recombination as the source of variability from which environmental events naturally select the fittest, such ‘natural selection’ constituting the cause of evolution. Recent biology increasingly challenges this view by casting genes as followers and awarding the leading role in the genesis of adaptations to the agency and plasticity of developing phenotypes—making natural selection a consequence of other causal processes. Both views of natural selection claim to capture the core of Darwin’s arguments in On the Origin of Species . Today, historians largely concur with the MS’s reading of Origin as a book aimed to prove natural selection the cause ( vera causa ) of adaptive change. This paper finds the evidence for that conclusion wanting. I undertake to examine the context and meaning of all Darwin’s known uses of the phrase vera causa , documenting in particular Darwin’s resistance to the pressure to prove natural selection a vera causa in letters written early in 1860 . His resistance underlines the logical dependence of natural selection, an unobservable phenomenon, on the causal processes producing the observable events captured by the laws of inheritance, variation, and the struggle for existence, established in Chapters 1–3 of Origin .

Since the synthesis of Darwinism and Mendelian genetics in the 1930s and 1940s, Darwinians have not regarded the struggle for existence as a cause of natural selection. (Radick, 2009 , p. 162). They admit variation as a  vera causa  in one case, they arbitrarily reject it in another, without assigning any distinction in the two cases. The day will come when this will be given as a curious illustration of the blindness of preconceived opinion. (Darwin, 1859a , p. 423).

Introduction

Before the seminal marriage of evolutionary theory with modern genetics, Gregor Mendel was commonly thought non-Darwinian—because the effects of genetic mutations were held to be discontinuous, and so incompatible with Darwin’s evolutionary dictum that ‘nature does not make jumps’ (e.g. Bateson, 1909 ; cf. Darwin, 1859a , pp. 171ff: ‘natura non facit saltum’). As soon as the ‘modern synthesis’ (MS; Huxley, 1942 ) gained influence over evolutionary science, its deployment of population genetics meant Mendel was recast as a Darwinian (Fisher, 1936 ; Sapp, 1990 ).

Like transformations have moulded understandings of Charles Darwin’s own work. Before the 1930s, his writings had played divers roles across different branches of science: biology; genetics; geology; biometrics; and taxonomy amongst them (e.g. Depew & Weber, 1994 ; Gayon, 1992 ). Come the MS, however, and Darwin got recast as the purveyor of a single idea—‘the best idea anyone has ever had’—meaning that, by century’s end, Darwinism had become synonymous with a belief in natural selection which was, purportedly, ‘the fundamental mechanism responsible’ for evolution (e.g. Dennett, 1995 , pp. 21, 46). As a result, Darwin’s many other observations and theses about how evolution worked were side-lined as wrong or irrelevant to contemporary science. At the same time, the MS endowed Darwin’s treatment of natural selection with a newly-narrowed, and still-dominant identity (Provine, 1988 ; Smocovitis, 1992 ). It became the cause or mechanism of evolution: chance environmental events blindly winnowing random genetically-caused variations in organisms’ DNA, so that descendant gene pools (and, only consequently , organisms) grow better adapted to their conditions of life than were their ancestor populations.

Syncing nicely with the MS, by the year 2000 historians of science had firmly established an MS-consistent reading of On the Origin of Species (Darwin, 1859a ; henceforth ‘ Origin ’) as a book primarily aimed to prove that ‘natural selection’ is the causal mechanism of evolution (e.g. Hodge, 2013 ; Hull, 2003 ; Pence, 2018 ; Ruse, 2005 ; Waters, 2003 ). Yet, over recent decades, biology’s MS has increasingly been challenged, qualified, or ‘extended’ by the findings of evolutionary science (e.g. Laland et al., 2015 ; Oyama et al., 2001 ; Walsh, 2015 ; West-Eberhard, 2003 ). In the process, scientists’ views have begun to diverge about the central thesis of Origin and, in particular, about how Darwin understood ‘natural selection.’ Such divergence presents a challenge to historians’ readings of Origin as purveying a view of evolution consonant with the MS. Does Darwin’s masterwork genuinely—but wrongly, according to a growing number of twenty-first century evolutionary biologists—equate natural selection to a ‘mindless, purposeless, mechanical process,’ as both modern synthesisers and their critics continue to claim (e.g. Dennett, 1995 , p. 34; Lewontin, 1983 , p. 275; Pigliucci et al., 2010 , p. 11)? In which case evolutionary biology is en route to becoming non -Darwinian. Or do historians’ MS-consistent readings of Origin miscast its arguments?

This essay tackles those questions. It aims to reassess historical evidence about Darwin’s take on the argument that natural selection provides the causal mechanism for evolutionary change. I will examine in detail: how Darwin himself used the phrase ‘ vera causa ’; the way Origin constructs its argument about the causes of evolution; and how Darwin defended that book against criticisms of his approach to scientific investigation. I also review the grounds for modern historiographic conclusions that Origin argues natural selection to be a causal mechanism. In this, I draw out conflicting uses of that ambiguous phrase ‘ verae causae ’ by Darwin scholars. I conclude by showing that understanding natural selection as an effect of other processes—not a cause in its own right—has critical significance for contemporary evolutionary theory. To that end, I will start my argument by briefly outlining the place of understandings of Darwin’s work in debates about the explanatory status of natural selection in today’s evolutionary science.

The contemporary scientific debate about natural selection

Twenty-first century science poses three kinds of challenge to MS understandings of evolution, under the banners: ‘evo-devo’; ‘developmental systems theory’ (DST); and a phenotype-first theory of adaptive change as led by ‘developmental plasticity.’ Despite their considerable differences (Bradley, 2020 , pp. 97–101), these new approaches have together been dubbed the ‘extended evolutionary synthesis.’ But they do not all challenge the idea of natural selection as cause (e.g. Laland et al., 2013 ; Pigliucci et al., 2010 ). A subset of these approaches, which do consistently challenge the causal interpretation of natural selection, have been dubbed, ‘developmental,’ ‘situated’ and ‘active’ Darwinism’ (Noble, 2020 ; Walsh, 2012 , 2015 ).

Evolutionary developmental biology (‘evo-devo’) addresses ‘the profound neglect of development in the standard modern synthesis framework of evolutionary theory’ (Muller, 2007 , p. 943). It seeks to understand both how developmental processes have evolved and how they may have helped cause the evolution of adaptations (Arthur, 2002 ). But evo-devo typically retains what its advocates call ‘Darwin’s’ conception of natural selection: as a causal ‘process’ or ‘mechanism’ which ‘acts’ at the population level (e.g. Arthur, 2002 , pp. 759–762; cf. Hall, 2012 , p. 187; Muller, 2007 , p. 94). DST typically theorizes phenotypic adaptation as developed via a system which incorporates an organism’s or population’s genes along with their ‘environments’—the term ‘environments’ including the ‘internal’ intra-cellular environment’s control of gene expression (epigenetics), plus developmental processes, alongside stable features of the ‘external’ environment (Oyama et al., 2001 , pp. 1–11). However, DST still confusingly casts natural selection both as an ‘emergent phenomenon’ (a higher-order effect?) stemming from ‘lower-level’ processes, and as a productive cause of adaptation (Griffiths & Gray, 2001 , p. 214; Weber & Depew, 2001 , pp. 244–249). Finally, West-Eberhard ( 2003 , pp. 33ff; Walsh, 2010 ) has advanced a comprehensive, evolutionary theory of the phenotype which conceptualises phenotypic development in terms of ‘plasticity’: the ability of an organism to react to an internal or external environmental input with reversible or irreversible changes ‘in form, state, movement, or rate of activity.’ West-Eberhard’s understanding of plasticity incorporates a stress on the agency of whole organisms, a theme that has become increasingly prominent over recent years (e.g. Bradley, 2020 ; Nicholson, 2014 ; Nicholson & Dupré, 2018 ; Noble, 2020 ; Walsh, 2015 ). She argues agentic plasticity ‘leads’ evolutionary change, with genes acting as ‘followers’ which subsequently stabilise adaptive phenotypic changes (West-Eberhard, 2003 , pp. 157–158). Her analysis makes natural selection an effect of other causes.

These new approaches expose a rift between two competing views of evolution by natural selection: the eighty-year-old gene-stressing MS and its derivatives (e.g. Dennett, 1995 ; Huxley, 1942 ), versus those contemporary views which stress the plasticity of organismic agency, and of phenotypic development, as what guides adaptation. Yet both these rivals claim direct descent from Origin . So we find Dawkins ( 2006 , p. xv, my italics) maintaining that his MS-based selfish-gene view of evolution ‘ is Darwin's theory , expressed in a way that Darwin did not choose but whose aptness, I should like to think, he would instantly have recognized and delighted in.’ Meanwhile, the new ‘developmental’ paradigm of evolutionary biology, writes Walsh ( 2010 , p. 336), ‘preserves more of the core of the Origin of Species than Modern Synthesis Replicator theory does’ (see too West-Eberhard, 2003 , pp. 186–193; 2008 ). In particular, Walsh ( 2012 , p. 192, my italics; drawing on West-Eberhard, 2003 ) argues that a crucial difference between these two Darwinisms is that, while the MS holds natural selection to be a causal force in its own right, developmental or active Darwinism cleaves closer to the Origin in making natural selection— not a cause of evolution, as per the MS view—but ‘a higher-order effect ’ of a number of other causal processes, most notably, of the struggle for existence and of individual development.

Which poses a question: which of today’s two Darwinisms better captures what Origin argues?

Origin ’s framing of natural selection and Darwin’s responses to its critics

The explanatory status of natural selection in Origin does not just concern today’s historians. It worried Darwin himself, particularly in the few months following his book’s launch. Several reviewers—including Darwin’s allies Charles Lyell ( 1859 ), Thomas Huxley ( 1860a , 1860b ) and Asa Gray ( 1860 )—had quickly queried the scientific orthodoxy of the book’s method of argument, and the comprehensiveness of its evidence for its conclusions. Darwin took pains to address these worries, not just in private letters, but in amendments to later editions of Origin , and published defences of its arguments.

In the first edition of Origin , Darwin ( 1859a , pp. 84, 146) habitually wrote as if natural selection were an intelligent agent, ‘intently watching each slight alteration’ in an organism’s structure and habits, so that it could ‘pick out with unerring skill each improvement’:

It may be said that natural selection is daily and hourly scrutinising, throughout the world, every variation, even the slightest; rejecting that which is bad, preserving and adding up all that is good; silently and insensibly working, whenever and wherever opportunity offers, at the improvement of each organic being in relation to its organic and inorganic conditions of life.

Writing in this vein, Darwin ( 1859a , pp. 85, 156) cast natural selection as a ‘power,’ which ‘acts by life and death,’ and so ‘causes’ extinction, for example. Not only antagonists (like Adam Sedgwick, 1859), but even allies like Charles Lyell (1860a) and Joseph Hooker ( 1860 ) complained Darwin had cast natural selection as a power akin to a deity, a ‘deus ex machina’ as Hooker ( 1862 ) later put it. Darwin denied the claim. (And later editions of Origin qualified his use of anthropomorphic language.) 1 Yet both the rhetorical organisation of his argument, and the fact that his book used an ordinary language immanently ‘imbued with intentionality,’ weakened these denials (Beer, 2000 , p. 81).

Darwin had launched Origin ’s argument with an account of ‘variation under domestication’ which celebrated the considerable changes in the forms of domesticated plants and animals (especially pigeons) effected by breeders’ powers of ‘artificial selection.’ He then extended this analogy to provide the framework for his exposition of ‘natural selection.’ In so doing, he rhetorically projected an image of ‘nature’s power of selection’ in the form of a human skill, though one of ‘far higher workmanship’ and producing modifications ‘infinitely better adapted to the most complex conditions of life’ than the ‘feeble’ ‘artificial’ efforts of stud-farmers and horticulturalists (Darwin, 1859a , pp. 84, 109). At the same time, when pushed, he strongly resisted the idea that ‘natural selection’ was an anthropomorphic causal agency (Young, 1971 ).

This contradiction arose because, as Beer ( 2000 , pp. xviii, 3, 48) put it, Origin ’s central argument ran directly ‘against the grain of the language available’ to Darwin as a Victorian man of science, a language which spot-lit ‘design and creation.’ Darwin’s struggle was to portray natural law and the uniformity of nature as things opposed to design and divine creation. Thus, viewed in the round—and despite the vagaries of the metaphorical way he had expressed his thesis—his book aimed to convince readers that natural selection was ‘ one general law , leading to the advancement of all organic beings, namely, multiply, vary, let the strongest live and the weakest die’ (Darwin, 1859a , p. 244, my italics; cf. Darwin, 1874 , pp. 48, 613). In 1861, he hastened to underline this aim by adding a caveat to the third edition of Origin ( 1861a , p. 85, my italics):

Several writers have misapprehended or objected to the term Natural Selection. Some have even imagined that natural selection induces variability, whereas it implies only the preservation of such variations as occur and are beneficial to the being under its conditions of life … It has been said that I speak of natural selection as an active power or Deity; but who objects to an author speaking of the attraction of gravity as ruling the movements of the planets? Every one knows what is meant and is implied by such metaphorical expressions; and they are almost necessary for brevity. So again it is difficult to avoid personifying the word Nature; but I mean by Nature, only the aggregate action and product of many natural laws, and by laws the sequence of events as ascertained by us . With a little familiarity such superficial objections will be forgotten.

Natural selection, Darwin here re-asserts, describes a ‘sequence of events as ascertained by us.’ It does not actively produce variations: it results from (the preservation of) variations—such preservation being something which itself ‘ results from the struggle for existence’ (Darwin, 1859a , pp. 5, 433, my italics).

This reading of Origin is reinforced by Darwin’s response to a fierce debate over the explanatory status of ‘natural selection’ in the early months of his book’s life. Origin itself labels natural selection in several ways, most commonly as a principle, a law, or a theory. Within weeks of its appearance, however, both private correspondents and public reviews opened a debate about the causal efficacy of natural selection. Had Darwin’s book proved natural selection to be a true cause (or ‘ vera causa ’) of the origination of species? His friends Huxley ( 1860a , 1860b ) and Gray ( 1860 ), while both hugely appreciative of Origin , concluded that Darwin had failed to prove natural selection was the effective cause of evolution. Others were more contemptuous. Palaeontologist Richard Owen ( 1860 ) pronounced natural selection a hypothesis resting on ‘a purely conjectural basis.’ Philosopher of science John Herschel dismissed Origin as ‘the law of higgledy-piggledy’ (Darwin, 1859b ; Hull, 2003 , pp. 181–182). And Darwin’s old geological mentor Adam Sedgwick ( 1860 , p. 335) found in Origin a ‘baseless theory.’

Darwin ( 1863a ) reacted to these criticisms by pointing out in a letter to the Athenaeum (provoked by Owen, 1863 ) that, though he continued to believe that ‘the theory, or hypothesis, or guess, if the reviewer likes so to call it, of natural selection’ provided the best explanation for the origin of species, such explanation ‘signifies extremely little in comparison with the admission that species have descended from other species and have not been created immutable.’ In short, the chief aim of his book was to win scientific acceptance of the fact that all species of creature were bound together by a web of affinities that comprised a ‘community of descent.’ His exposition of natural selection was first and foremost, aimed to gain assent for the fact of evolution. Its exact explanatory status was of secondary importance.

Meanwhile, behind the scenes, Darwin worked hard to garner evidence which would satisfy Huxley’s doubts about the explanatory status of natural selection. Huxley ( 1860b , pp. 74–75) did not himself use the term ‘ vera causa ’ in his review of Origin . His argument was that Darwin’s thesis on natural selection would remain ‘a hypothesis,’ and not yet a ‘theory of species,’ until ‘positive evidence’ could be produced that a group of animals (or plants) had, ‘by variation and selective breeding, given rise to another group which was, even in the least degree, infertile with the first.’ Besides conducting many (unsuccessful) experiments of his own to prove the sterility of inter-breed hybrids in flowers, Darwin recruited a host of zoologists, botanists and horticulturalists to find evidence to fill the gap that Huxley had identified, including Hooker, Muller, Tegetmeier, Gray and many others—even sending an open letter to the readers of the Journal of Horticulture to beg for relevant facts (Darwin, 1862 ). The chapter on hybridism in the fourth edition of Origin (Darwin, 1866 ) was expanded to discuss the most promising new findings, with the hope of satisfying Huxley. The chapter on hybridism in The Variation of Animals and Plants under Domestication (Darwin, 1868 ) also aimed to answer Huxley’s criticism. And later, the opening paragraphs of the last chapter in the final edition of Origin (Darwin, 1876 ) were enlarged to address the sterility of hybrids. Huxley remained unconvinced.

But Darwin’s strongest and most immediate response to critics of the scientific status of Origin ’s claims was directly to contest the need to prove natural selection a vera causa . Within three months of the book’s publication, he told his closest friend Hooker (Darwin, 1860a ) that Huxley ‘rates higher than I do the necessity of Natural Selection being shown to be a vera causa always in action.’

Darwin contests the need to prove natural selection a vera causa

Three kinds of consideration underlined the subordinate importance of the vera causa criterion for Darwin. Firstly, the exact nature of the standard of proof for a vera causa had been disputed so much over previous decades that, by 1859, its meaning was extremely loose (Ruse, 1975 ). Origin (Darwin, 1859a , p. 423) lampoons the resultant imprecision in scientific identifications of natural phenomena as verae causae : while ‘several eminent naturalists … admit variation as a  vera causa  in one case, they arbitrarily reject it in another, without assigning any distinction in the two cases’—such arbitrary identifications revealing only the power of ‘the blindness of preconceived opinion.’

The inconsistency of scientific judgements about verae causae that Darwin disparaged is confirmed by three trail-blazing historiographical essays (published in the 1970s) which concluded that Origin aimed to prove natural selection a vera causa , in that all three disagree about what Darwin would need to have done to achieve this aim (Hodge, 1977 , 1989 ; Hull, 1973 , 2003 ; Ruse, 1975 , 1999 , p. 57). This disagreement continues today, as Sect.  4 of this paper documents. One historian, Greg Radick ( 2002 , p. 13, my italics), has even glossed Darwin’s adherence to the vera causa ideal as meaning that Origin sought to show that ‘ the causes that together produced natural selection —variation, inheritance and the struggle for existence—were all “true causes,” that is, causes evidenced independently of the facts they were held to explain.’ Radick’s reading—that Origin frames natural selection as the consequence of several other, directly-observable (and hence ‘true’) causal processes—points directly to the argument I advance here. But it stands in stark conflict with the views of the Origin ’s strategy elaborated by Hodge and Ruse in the 1970s, as well as many more recent historiographic claims.

Secondly, Darwin’s own most-repeated criterion for the scientific reality of natural selection was that it could explain the several distinct ‘large classes of facts’ that Origin ( 1876 , p. 568) argued natural selection did explain. This standard of proof was akin to what William Whewell ( 1840 ) called a ‘consilience of inductions’: ‘the best kind of science … comes when different areas of science are brought together and shown to spring from the same principles’ (Ruse, 1975 , p. 163). Yet the status of such consilience vis-à-vis verae causae remains uncertain. According to some historians, such ‘consilience’ was a hallmark of verae causae (Ruse, 1975 , 1999 , p. 58; Waters, 2003 ). Others, like Hodge ( 1989 , pp. 171–173, my italics) argue that Whewell ‘offered his consilience ideal as an alternative ’ to the vera causa ideal.

The ‘large classes of fact’ or ‘different areas of science’ which Origin ( 1859a , pp. 415, 420; 1876 , pp. 137, 424) treats as explained by ‘the same principle’ of natural selection include: the homologous forms of rudimentary, embryological, and anatomical structures in taxonomically related species (e.g. wing of the bat, fin of the porpoise, leg of the horse, human hand); the fact that pre-evolutionary taxonomic classification could be arranged within ‘a few great classes, in groups subordinate to groups, and with the extinct groups often falling in between the recent groups’; endemic species on oceanic islands being related to the nearest source of immigrant species (as in the Galapagos archipelago); the gradual diffusion of dominant forms in the geological record; the co-adaptations of different species to each other within the same habitat; and the lack of perfection of some adaptations—‘the sting of the bee, when used against an enemy, causing the bee’s own death,’ ‘drones being produced in such great numbers for one single act, and being then slaughtered by their sterile sisters,’ ‘the astonishing waste of pollen by our fir-trees’ (Darwin, 1876 , pp. 415, 419).

Darwin reverted to his consilient criterion of proof time and again, not just in Origin , but in his other books (e.g. Darwin, 1874 , p. 24; 1890 , p. 113), and in his letters. For example: ‘It seems to me that an hypothesis is developed into a theory solely by explaining an ample lot of facts’ (Darwin, 1860b ). In contrast, Darwin’s publications never refer to natural selection as a vera causa (nor do they ever refer to it as a mechanism; Ruse, 2005 ). In fact Origin was the only one of his books to reference verae causae at all. It uses the phrase thrice: once to refer to ‘community of descent’; once to suggest that, when a single species occurred at ‘several distant and isolated points,’ the ‘the vera causa of ordinary generation with subsequent migration’ was a better explanation for it doing so than the ‘miracle’ of several separate divine creations; and once, as we just saw, to ridicule the arbitrariness of the assignment to ‘variation’ of the status of a vera causa in the creationist arguments of ‘several eminent naturalists’ (Darwin, 1859a , pp. 159, 352, 482). These three usages all occur in polemics directed at creationists—the last stating directly that naturalists’ identification of verae causae owed more to prejudice than to science. Which might suggest the phrase functioned at best rhetorically in Origin , which variously enlisted, and questioned, its gravitas as a shibboleth of scientific proof.

Thirdly, three months after Origin came out, Darwin discovered a powerful parallel between natural selection and Newton’s law of gravity. In February 1860, Darwin was reading David Brewster’s ( 1855 ) Memoirs of Sir Isaac Newton . This was the month he was most acutely focused on arguments, like Huxley’s ( 1860a ), that Origin had failed to prove natural selection causally efficacious. Brewster ( 1855 , pp. 282ff) recounted how, in 1710, Leibnitz had attacked Newton’s theory of gravity as ‘introducing occult qualities and miracles into philosophy.’ Newton retorted that the theory of gravity was:

proved by mathematical demonstration, grounded upon experiments and the phenomena of nature; and that to understand the motions of the planets under the influence of gravity, without knowing the cause of gravity , is as good a progress in philosophy as to understand the [movements of the clockwork of a clock, as a clockmaker does] without knowing the cause of the gravity of the weight which moves the machine … (Newton, 1711, quoted in Brewster, 1855 , p. 283, my italics)

Darwin pounced on this passage because it underlined a distinction between law and cause, as Darwin swiftly pointed out to Lyell and Gray. Darwin’s ( 1860c ) comment to Lyell was that, though Leibnitz held the law of gravity to be unscientific, mysterious or ‘occult’ (because gravity had not been directly observed), Newton’s law nonetheless added to our knowledge because it explained ‘the movement of wheels of clock, though the cause of descent of the weight could not be explained ,’ adding to Gray the next day (Darwin, 1860d ): ‘This seems to me rather to bear on what you say of Nat. selection not being proved as a vera causa.’

What interested Darwin about Newton’s reply to Leibnitz was that the law of gravity brings under one descriptive formula various ‘sequences of event as ascertained by us’—tidal flows, falling apples, clockwork, and planetary orbits—even though Newton could not say what caused those events. 2 Likewise, the origin of new species by natural selection was too slow to be observed. Yet, like Newton’s law, Darwin’s argument gave coherence to various ‘sequences of event’—palaeontological succession of types, geographical distribution, taxonomic nesting, homologies of anatomical structures in related taxonomic classes, in rudimentary organs and in embryos etc.—however much what caused those events remained open to question.

In the same month that he was reading Brewster, Darwin ( 1860e ) developed a second parallel with physics, this time between his ‘hypothesis’ of natural selection and the wave or ‘undulatory’ theory of light, which, by the 1850s, was becoming increasingly favoured over Newton’s ( 1704 ) corpuscular theory. Of course, no one had ever observed the undulations in the so-called ‘luminiferous ether’ which constituted light, according to Robert Hooke ( 1664 ) and Christiaan Huygens (1690). Yet, said Darwin, the wave theory ‘groups together & explains a multitude of phenomena,’ such as the interference patterns seen in Thomas Young’s ( 1804 ) diffraction experiment, and so was ‘universally now admitted as the true theory.’ ‘The undulatory theory of Light is very far from a vera causa ,’ noted Darwin ( 1860f , my italics), yet it was scientifically ‘allowable (& a great step) to invent the undulatory theory of Light.’ So why should not scientific procedure allow Darwin ( 1860g ) also to ‘invent [the] hypothesis of natural selection … & try whether this hypothesis … does not explain (as I think it does) a large number of facts in geographical distribution—geological succession—classification—morphology, embryology &c. &c.’? 3 By implication, Darwin is here acknowledging that natural selection, while having scientific value, is also ‘very far from a vera causa.’

Darwin’s parallel between the law of gravity and natural selection was swiftly spliced into the last chapter of the third edition of Origin ( 1861a , pp. 514–515), which thenceforth noted that, though ‘the law of the attraction of gravity’ had been attacked by Leibnitz because no one knew ‘what is the essence of the attraction of gravity,’ yet ‘no one now objects to following out the results consequent on this unknown element of attraction.’ Origin ’s final edition saw a further inclusion—Darwin’s ( 1876 , p. 421) parallel between natural selection and the wave theory of light—a parallel which had already been developed at greater length in the exposition of natural selection opening Variation (Darwin, 1868 , vol.1, pp. 8–9). 4

Modern vera causa readings of Origin

I now consider how my reading of Origin bears on evidence for contemporary historians’ conclusion that the main aim of Origin is to prove natural selection has been the ‘true cause’ ( vera causa ), ‘mechanism’ or ‘causal force’ effecting the origin of species (e.g. Gildenhuys, 2004 ; Hodge, 2013 ; Pence, 2018 ; Ruse, 2005 ). It should be remembered that, whilst vera causa interpretations of Origin are widely assumed by today’s Darwin scholars, they disagree amongst themselves as to what vera causa might have meant to Darwin in 1859. Pence ( 2018 ) finds seven current historiographic interpretations of the philosophy of science underpinning Origin —and his list is not exhaustive.

Neither Darwin nor Origin ever claim that natural selection is a vera causa —though, as we saw above, Origin does use this phrase in connection with three other facets of Darwin’s argument. Nor does Darwin anywhere, in his publications, notebooks or private correspondence, say that the book was designed to prove natural selection is a vera causa . On the contrary, he said the book was designed to prove that all species share in a community of descent (besides which, he added, the validity of any claims he had made about natural selection signified ‘extremely little’; Darwin, 1863a , see Sect.  3 ).

So: what evidence do today’s historians advance to back their contention that Darwin wrote Origin to prove natural selection a vera causa ? They largely ignore the evidence I reviewed in Sect.  3.1 , quoting instead the first third of the postscript to a letter Darwin wrote to botanist George Bentham, in May 1863. This letter concerned an address, intended to support Origin , which Bentham was preparing to give to the Linnaean Society (of which he was president) in two days’ time. Bentham (1863, my italics) was worried that Darwin’s theory could not explain why some species of ‘northern hemisphere’ plants were found in Tasmania and Australia’s Victorian Alps to ‘have gone through so many thousand generations in both hemispheres unaltered ,’ whilst other species of such plants had changed so much as to become almost unrecognisable. The postscript to Darwin’s ( 1863b ) reply reads as a kind of executive summary to help the doubting Bentham prepare his imminent address by clarifying three alternative kinds of grounds upon which a belief in natural selection could be based:

In fact the belief in natural selection must at present be grounded entirely on general considerations. (1) on its being a vera causa, from the struggle for existence; & the certain geological fact that species do somehow change (2) from the analogy of change under domestication by man’s selection. (3) & chiefly from this view connecting under an intelligible point of view a host of facts.— When we descend to details, we can prove that no one species has changed: nor can we prove that the supposed changes are beneficial which is the groundwork of the theory. Nor can we explain why some species have changed & others have not … 5

A century later, in 1975, Ruse concluded that this postscript shows that, useful though Darwin believed the analogy between artificial and natural selection might be (Darwin’s 2nd point), ‘the chief proof for Darwin of the truth of his theory was that it had explanatory power in all of these many diverse areas’ (Darwin’s 3rd point). In a footnote to the same article, Ruse ( 1975 , p. 177) challenged the import of the postscript’s first point, saying that, although Darwin here ‘wrote of natural selection as a vera causa ,’ by 1859 this term ‘was used almost as loosely as “deduction”.’ Elsewhere, however, both in his 1975 article and in later works, Ruse ( 1975 , p. 175; 1999 ; 2005) agrees with Hodge ( 1977 , p. 238) that in Origin , Darwin was ‘committed’ or ‘desperately keen’ to show that ‘his evolutionary reasonings were based on a vera causa , natural selection.’ Yet these two historians disagree as to whether Darwin’s ‘commitment’ was informed by John Herschel’s interpretation of Thomas Reid’s understanding of vera causa (Hodge, 1977 , 1989 ), or by the incompatible views (according to Hodge, 1989 , p. 172) of William Whewell (Ruse, 1975 , 1999 ).

Hodge ( 1977 , pp. 240–241; Hodge, 1989 , p. 190; 2013, p. 2273) also disagrees with Ruse about Darwin’s postscript to Bentham. He reads its three points as a rationale for decoding Origin ’s entire structure as something rooted in a three-step strategy to prove natural selection is a vera causa (a strategy which I will describe shortly). Gildenhuys ( 2004 , pp. 594, 605) and Pence ( 2018 ) also ground their arguments on the postscript (though both disagree with Hodge’s reading). None of these articles by Ruse, Hodge, Gildenhuys, or Pence, which cite the postscript to Bentham, discusses the far more substantial correspondence Darwin had had in the opening months of 1860, regarding the causal status of natural selection (discussed in Sect.  3.1 ). Nor do they reference the changes Darwin made to the third and later editions of Origin , reflecting the lasting importance of the points made in that correspondence.

Whether in his publications, his notes or his correspondence, Darwin rarely mentioned verae causae. His only published mention of a vera causa prior to its appearance in Origin was twenty years earlier, in his geological ‘Observations on the Parallel Roads of Glen Roy’ (Darwin, 1839 ; see Sect.  5.1 below). This paper had over-confidently (and falsely) identified the action of river deltas flowing into the sea as a vera causa for the formation of the ‘buttresses’ found below Glen Roy’s ‘parallel roads,’ a deduction Darwin later accounted ‘a great failure, and I am ashamed of it’ (Darwin, 1958 , p. 84). Bar Origin , none of his other books or papers mention verae causae. The sixty plus years of his vast correspondence contain just nine mentions of verae causae. Seven of these occur in the vera causa debate about natural selection during the five months after the book came out—all of which dispute ‘the necessity of Natural Selection being shown to be a vera causa always in action’ (Darwin, 1860a ; see Sect.  3.1 ).

Darwin’s last ever use of the term vera causa is in the oft-cited postscript to Bentham. This comes more than three years after his earlier flurry of correspondence about the causal status of natural selection. By May 1863, Darwin ( 1861a , 1861b ) had re-edited the text of Origin to underline his distinction between cause and law, and now believed that, among his scientific allies, there were many who had accepted natural selection was a vera causa (whatever that meant), including John Stuart Mill. 6 He also knew that many other eminent men of science (including Huxley, Sedgwick, Owen, and Whewell) continued to deny it this status. So, when he noted to Bentham (a worried ally) that the belief in natural selection ‘must at present be grounded entirely on general considerations,’ he did not , in so many words, claim that natural selection was a vera causa . Because, as his whole letter underlined—and the much-quoted postscript reiterates 7 —he knew that there could be no direct observational evidence (‘details’) that even ‘one species has changed.’ Even for those who believed natural selection to be a vera causa , its status as such could only be deduced from things that could be observed (which, according to some interpretations of the concept, disqualified it as a vera causa , because, in the words of one critic, his theory was ‘not inductive —not based on a series of acknowledged facts’; Sedgwick, 1860 , p. 334).

The postscript’s first point therefore acknowledges that, at best, natural selection’s status as a vera causa could only be derived indirectly from observational evidence for ‘the struggle for existence; & the certain geological fact that species do somehow change.’ It was for this reason that—though the analogy between artificial and natural selection provided another possible rationale—the postscript went on to stress that, for Darwin , the chief basis for a belief in natural selection was not that it was a vera causa , but consilience : its ‘connecting under an intelligible point of view a host of facts’ (see Sect.  3.1 ).

Claims that Origin is structured to prove natural selection a vera causa are further weakened by Hodge’s (e.g. 1977 , p. 239; 1992 ; 2013 ) own efforts to force the book into the three-step framework he deems such a proof should take: ‘in explaining any phenomenon, one should invoke only causes whose existence and competence to produce such an effect can be known independently of their putative responsibility for that phenomenon.’ This implies, says Hodge, that Origin should comprise three sections, each section containing a group of chapters evidencing in turn: the existence of natural selection; the competence of natural selection to produce species-change; and, finally, evidence that natural selection really had been responsible for species-change. ‘Unfortunately,’ says Hodge ( 1977 , pp. 238, 242), Origin ‘violates’ this structure, ‘misleadingly’ making ‘successive departures’ from it—departures which ‘were eventually enough to render the strategy and organization of his most famous book unhelpfully and quite unnecessarily obscure.’Which means the three ‘general evidential considerations’ upon which Darwin should have focused, ‘ do not map onto the Origin ’s three clusters of chapters’ (Hodge, 1977 , p. 244; 2013 , p. 2274, my italics).

Origin on what effects natural selection

Modern debates about Darwin’s putative ‘commitment’ to the vera causa principle and his associated ‘epistemological self-consciousness’ (e.g. Hodge, 1977 , p. 238; 2000, p. 29) attain a high degree of philosophical sophistication—far higher than any discussion to be found in Darwin’s own writings. Historiographers and philosophers of science make superfine distinctions between the epistemologies that are deduced to have influenced, or not to have influenced, Darwin’s authorial consciousness. Against this, Hull’s ( 1973 , 2003 ) essays repeatedly demonstrate how shallow an understanding of epistemological issues—and the arguments of contemporary philosophers of science like Mill—Darwin (and Huxley) actually possessed.

As Darwin’s autobiography (1958, p. 140) candidly admitted, ‘my power to follow a long and purely abstract train of thought is very limited.’ And while he did pay some attention to metaphysical subjects in his twenties, his attitude to the topic had become increasingly jaundiced as his reading progressed. During 1838, he spent several months absorbing the opinions of Hume, Mackintosh, Abercrombie, Comte and Ferrier, all of whom belittled metaphysics as, for example, ‘a name of reproach and derision’ (Mackintosh, 1830 , p. 4). By October of that year, he had concluded that: ‘To study Metaphysics, as they have always been studied appears to me to be like puzzling at astronomy without mechanics … we must bring some stable foundation to argue from’ (Darwin, 1838 , p. 5). In later years his use of the term ‘metaphysical’ became derogatory. 8 Books or articles that his letters dubbed ‘metaphysical,’ were condemned as ‘mere verbiage,’ being ‘barely intelligible,’ dealing in ‘far-fetched analogies,’ and constituting ‘rubbish’ produced by a ‘wind-bag’ with ‘muddled... brains’ and ‘an entire want of common sense’ (Darwin, 1845 , 1857 , 1860h , 1861b , 1864 , 1871 , 1874 , p. 78).

Given the improbability that Darwin adhered to a refined philosophical understanding of what vera causa meant when he used the term, I now try to decode what it did mean to him. First, by emphasising the most obvious common denominator in the various conflicting philosophical understandings of verae causae current in his day. And second, by examining what Darwin assumed on the few occasions when he himself did mention verae causae .

Despite a slim evidential basis, modern Darwin scholars have imaginatively constructed several contrasting pedigrees for Darwin’s understanding of verae causae —whether via Thomas Reid (Hodge, 1989 , p. 171), William Whewell (Ruse, 1975 ), John Herschel (Gildenhuys, 2004 ; Pence, 2018 ), or Charles Lyell (Rudwick, 2005 ; Sponsel, 2018 ). Whatever the merits of these different genealogies, all four hypothesised sources share one stress: the need for first-hand observational evidence in establishing the existence of a true cause. Reid insisted on ‘direct experiential acquaintance … as the only acceptable form of evidence for the known truth, reality or existence of a cause’ (Hodge, 1989 , p. 169). Whewell (like Sedgwick and Huxley) objected to Origin because no-one could ‘adduce a single example of one species evolving in nature into another. Nor had plant and animal breeders, through all their efforts, succeeded in producing a single new species’ (Hull, 2003 , p. 184). Herschel ( 1830 , #138) held that the best way of establishing a vera causa was from ‘experience [showing] us the manner in which one phenomenon depends on another in a great variety of cases.’ And Lyell’s uniformitarianism leant on the argument that causes which we can directly observe in the present , like the slow action of coastal waves, can be used to reconstruct the vast prehuman past as recorded in rock strata, so long as we assumed that ‘the same causes … had been at work with the same intensities and in the same overall circumstances’ from the time the first rock formed through to the most recent (Hodge, 2000 , pp. 28–29).

Darwin’s own usage of ‘ vera causa ’

Turn to Darwin’s own usage of ‘ vera causa ’ and we also find a stress on direct observation. His ill-fated Glen Roy paper explains the ‘buttresses’—flat-topped accumulations of alluvial gravel and other debris below the lowest of the parallel roads (see Fig.  1 ) —as left-over ‘deltas’ made by rivers or ‘streamlets.’ These streamlets he deemed to have formed the deltas or ‘raised beaches’ of the roads themselves, at the level which the streamlets flowed into the sea before the Glen was tectonically raised above present-day sea-levels by a sequence of crustal uplifts. (Darwin argued that the problematic lower-level buttresses must be remains of alluvial deposits made by the deltas of these same streamlets after further, less dramatic, crustal liftings of the land.) In this way, he exploited the easy-to-observe fact of the alluvial action of river deltas, which was already a central plank of previous geological explanations for the parallel roads (Rudwick, 1974 , pp. 106–107). Darwin’s paper (1839, p. 52) sums up this step in its argument by confidently asserting: ‘no one can doubt [t]hat this intervening cause [delta formation by rivers] has been … a  vera causa .’

Darwin’s ( 1839 ) illustration of the parallel roads of Glen Roy. The ‘buttresses’ are depicted by bent lines which represent bulging piles of rocky debris (e.g. below the lowest of the three roads)

His only other non- Origin -related use of the phrase was in a letter to William Redfield in February 1840. Redfield ( 1839 ) had just published an article relating ‘a few cases in which whirlwinds of great activity and violence appeared to have resulted from the action of fires.’ Darwin, who had long puzzled over the origin of waterspouts seen on his Beagle voyage, added an observation to Redfield’s list. This regarded a whirlwind and waterspouts resulting from an island-forming, submarine, volcanic eruption, observed by a Captain Tilliard off the Azores in June 1811. 9 Darwin ( 1840 ) wrote: ‘Taking your account of the whirlwinds produced by artificial fires, we here see the vera causa of one set of waterspouts.’

In Darwin’s response to Huxley’s critique of Origin , he also construes what he summarised as Huxley’s demand to prove natural selection a vera causa as a demand to produce what Huxley would recognise as ‘positive’ observational evidence (Sect.  3 ). Witness Darwin’s persistent efforts to produce such evidence from plant-experiments in his own garden, as well as from other horticulturalists and animal-breeders, to prove that domestic breeds had separated so far as to be ‘sometimes sterile with other breeds’ (e.g. Darwin, 1863b ; see Sect.  3 ).

It is this observation-based sense of evidence which informs the chapter-plan of Origin . Because, of course, as the latter two-thirds of the postscript to Bentham confirm, Origin ’s proposed ‘principle,’ ‘hypothesis,’ ‘theory’ or ‘general law’ of natural selection could never be observed to produce the detailed results Darwin’s book claimed that it had produced. The origination of new adaptations and new species was a process which the book held to take anything from ‘many thousands’ to ‘an almost infinite number’ of successive generations (Darwin, 1859a , pp. 114, 481). 10 Hence, as when Lyell’s Principles set out the geological processes observable in the here and now by which he would explain the formation of geological features dating from the earth’s remotest past (Rudwick, 1970 ), Origin sets out from what could be empirically witnessed in order to deduce what could not be witnessed.

Origin ’s presentation of natural selection

Origin ’s first three chapters elaborate several sets of empirical ‘laws,’ that is, statements based on repeated observations that describe (and thus predict) a ‘sequence’ of natural events: ‘laws of inheritance’ (Ch.1); laws of ‘variability’ and ‘correlated growth’ (Ch.2); and the law or ‘doctrine’ of Malthus, that populations of plants and animals have the reproductive capacity to increase at a ‘geometrical ratio,’ whilst food supplies, at best, increase at an arithmetical rate, resulting in a ‘struggle for existence’ (Ch.3). Each of these chapters details the many kinds of observable event which the said laws cover, giving copious examples. The laws themselves—whether of inheritance or variation—are, for ‘the most part unknown,’ or, at best, ‘dimly understood,’ Origin says (Darwin, 1876 , pp. 9–10). Yet, the existence of the phenomena these laws are meant to describe is hard to question, given the detailed observations and experiments Origin recounts regarding: the facts of inheritance in domesticated varieties of animal and plant; the multitudinous variability or ‘individual differences,’ whether in wild species, subspecies and varieties or in domestic breeds; and of the various kinds of competition and mutual aid (as in ‘social plants’) entailed in what Darwin underlined was a ‘metaphorical’ struggle between members of the same and different species to survive, thrive, and reproduce (Darwin, 1859a , pp. 62–63, 70–71).

Whilst inheritance and variability were necessary preconditions for natural selection, its principle engine was the struggle for life, which would winnow the more useful variations in a given habitat from the less. This struggle was a theoretical construct in Malthus. The aim of Origin ’s third chapter was to line up ‘better evidence on this subject than mere theoretical calculations, namely … numerous recorded cases,’ showing both species’ explosive potential for fecundity, and the vulnerability of individuals to a variety of environmental and inter-organism challenges (Darwin, 1859a , p. 64). Chapter Three particularly stresses the ‘web of complex relations’ between different creatures’ fates and those of the other organisms in their habitat (Darwin, 1859a , p. 73). Importantly, this metaphorical ‘struggle for life’ did not just betoken competition, but relative reproductive success and, the ‘dependence of one being on another,’ whether from different species (as with the symbiosis between moths and orchids), or from the same species, as with ‘social plants’ and ‘social animals’ who render ‘mutual aid’ to one another (Darwin, 1876 , p. 50; Bradley, 2020 ).

The logical dependence of Origin ’s fourth chapter, ‘Natural Selection’—and the book’s central thesis—on its first three chapters (on inheritance, variation, and the struggle for existence respectively) is reiterated throughout the book, from its first pages 11 :

As many more individuals of each species are born than can possibly survive; and as, consequently, there is a frequently recurring struggle for existence, it follows that any being, if it vary however slightly in any manner profitable to itself, under the complex and sometimes varying conditions of life, will have a better chance of surviving, and thus be naturally selected. From the strong principle of inheritance, any selected variety will tend to propagate its new and modified form (Darwin, 1859a , p. 5, my italics;).

Through exegesis of its central argument:

… it may be asked, how it is that varieties, which I have called incipient species, become ultimately converted into good and distinct species, which in most cases obviously differ from each other far more than do the varieties of the same species? How do those groups of species, which are called distinct genera, and which differ from each other more than do the species of the same genus, arise? All these results … follow inevitably from the struggle for life (Darwin, 1859a , p. 61, my italics)

To its last page:

It is interesting to contemplate a tangled bank, clothed with many plants of many kinds, with birds singing on the bushes, with various insects flitting about, and with worms crawling through the damp earth, and to reflect that these elaborately constructed forms, so different from each other, and dependent upon each other in so complex a manner, have all been produced by laws acting around us. These laws, taken in the largest sense, being Growth with reproduction; Inheritance which is almost implied by reproduction; Variability from the indirect and direct action of the conditions of life, and from use and disuse; a Ratio of Increase so high as to lead to a Struggle for Life, and as a consequence to Natural Selection , entailing Divergence of Character and the Extinction of less improved forms (Darwin, 1859a , pp. 489-490, my italics).

Such dependence is unavoidable, given the impossibility of gathering within a single human lifetime any eyewitness evidence for the efficacy of natural selection. This logic impels us to reject any claim that Origin makes ‘successive independent evidential cases … for natural selection existing at present ’ (e.g. Hodge, 2000 , p. 30). Such a statement would only make sense if natural selection were already assumed to be the consequence of the contributory laws copiously evidenced to ‘exist at present’ in the book’s opening three chapters—precisely the assumption writers like Hodge aim to overturn.

To underline this point, witness a crucial contrast between the first three chapters and the fourth. Unlike its predecessors, the supporting materials in Chapter Four ‘Natural Selection’ are not the fruit of first-hand observation, being presented entirely in the form of ‘ imaginary illustrations’ (Darwin, 1859a , p. 90, my italics; Bradley, 2011 ).

It is not until the latter parts of the fourth chapter that Origin starts seriously to discuss the causes for the laws evidenced in the book’s first three chapters— and thus of natural selection. The causal mechanism producing the laws of inheritance (i.e. the transmission of heritable characters) is bypassed. 12 Instead Chapters Four, Five (‘Laws of Variation’) and Six (‘Difficulties of the Theory’) proceed to explicate several causal processes that produce variations . Here, pride of place goes to the role of changed habits in directing selection. This theme stands out in Chapter Four’s illustrations of Darwin’s ‘principle of divergence’ of character (cf. adaptive radiation): Origin asks us to imagine the case of a carnivorous quadruped, ‘of which the number that can be supported in any country has long ago arrived at its full average’:

If its natural power of increase be allowed to act, it can succeed in increasing (the country not undergoing any change in conditions) only by its varying descendants seizing on places at present occupied by other animals: some of them, for instance, being enabled to feed on new kinds of prey, either dead or alive; some inhabiting new stations, climbing trees, frequenting water, and some perhaps becoming less carnivorous.

Darwin ( 1859a , p. 179; 1876 , p. 8) formalized this habit-first causal process for the evolution of transformative adaptations—as when ‘a land carnivorous animal’ had been ‘converted into one with aquatic habits’—in a way that showed how ‘changed habits produce an inherited effect.’ Origin ’s clearest example of this non-Lamarckian process highlighted how ‘transitional habits’ had plausibly resulted in the evolution of flying squirrels 13 :

Origin (Darwin, 1859a , pp. 179–186) asks us to imagine that, a long time ago, some adventurous, flightless squirrel-ancestors had formed a new habit of launching themselves, not just from branch to branch, but from tree-top to tree-top. Tree-surfing would put a new premium on glide-friendly changes to the squirrels’ physique (stronger spring at take-off, better depth vision, lighter body-weight, more aerodynamic tail, broader flanges of skin between front and back legs). Any chance heritable variation that fitted them better to their novel habit would have increased their reproductive success compared to unchanged conspecifics. Hence, ‘it would be easy for natural selection to fit the animal, by some modification of its structure, for its changed habits.’ Thus, while the production of what we now know as genetic variations—which must have stabilized the bodily changes that make tree-surfing easier for squirrels—might be random, the direction of adaptation would be set by the non-random agentic innovations of the ancestral squirrels.

‘Changed habits’ included ‘use and disuse,’ not just in animals but in plants. Of changed habits in plants, Darwin ( 1859a , pp. 139–143) cited the ‘acclimatisation’ of, for example, ‘the pines and rhododendrons, raised from seed collected by Dr. Hooker from trees growing at different heights on the Himalaya, [which] were found in this country to possess different constitutional powers of resisting cold’—seeds taken from higher in the mountains being found habitually more resistant to chilly British weather than their cousins from the mountains’ lower slopes. Darwin (e.g. 1859a , p. 76) typically framed the qualities of an organism in terms of ‘strength, habits, and constitution.’ The fate of variations in anatomy very often depend on an organism’s habits, according to his accounts, as, for example, with: the displays that feature sexual ornaments and the fights, which, he argued, must have led to the sexual selection of tusks and other weapons of sexual rivalry; the eating habits of birds with different shaped bills (e.g. finches in the Galapagos archipelago; Lindholm, 2015 ); insects’ adaptations to feeding from and so pollinating certain species of flower; closely-related animal species avoiding hybridisation by ‘haunting different stations’ of a given habitat; or the growth of hardness in pigeon chicks’ beaks (used for cracking their way out of their egg) (Darwin, 1859a , pp. 87, 103; 1882 ).

Other causes of variability proposed by Origin included ‘direct action of the environment’—on the ‘plastic’ (Darwin’s word: 1876 , pp. 62, 106, 438) quality of the reproductive system, and the creature’s ‘whole organisation.’ Such action depended both on the nature of the organism and the nature of the conditions (e.g. climate, altitude), the nature of the organism being ‘much the more important,’ according to Darwin ( 1876 , p. 6)—an emphasis now re-echoing through today’s post-MS biology with its so-called ‘return to the organism’ (e.g. Lewontin, 1983 ; Nicholson, 2014 ; Walsh, 2015 ).

Finally, the domain of phenotypic variability is not coextensive or neatly aligned with those ‘variations’ of relevance to a theory of natural selection. Not only may some of the ordinary doings of organisms fail to impinge on the struggle for existence—‘the war of nature is not incessant’ (Darwin, 1859a , p. 79). Even those that do so impinge may not result solely in ‘advantages,’ but also—as Chauncey Wright ( 1870 , p. 293) argued—‘limiting disadvantages,’ likely to undermine fitness. The Descent of Man (Darwin, 1874 , p. 571) proposed that Wright’s argument had ‘an important bearing on the acquisition by man of some of his mental characteristics’—citing in illustration how processes that (adaptively) ensured group cohesion, could simultaneously foster maladaptive customs and superstitions in some peoples. Examples included tribes where infanticide and cannibalism were customary, as reported by some ethnographers, plus, in Darwin’s own society, mating choices based on ‘mere wealth or rank’ (Darwin, 1874 , pp. 121–122, 617).

Given Darwin’s identification of verae causae with processes that can be directly observed, it makes sense that he should have structured Origin to prove the existence of natural selection—something unobservable —as being a higher-order consequence of other observable (causal) processes. To recognise Origin presents natural selection as an effect of other causes, not a cause in its own right, is not merely a matter of textual exegesis, however. Such recognition has dramatic repercussions for the contemporary interpretation of evolutionary theory: because it fells the central pillar of gene-based MS constructions of the natural world—the belief that evolution is caused by natural selection. This forces on evolutionary scientists the need to seek a brand new conceptualisation of the relationship between evolution and what Walsh ( 2015 , Ch.2.1) calls ‘the normal activities of organisms,’ including ourselves.

Here, the uptake of evolutionary theory by psychologists furnishes an apt illustration. Psychology is the central scientific site for examining the normal activities of organisms, particularly of human beings. So, how would adoption of the view that natural selection is ‘an analytic consequence’ (Walsh, 2015 , Ch.2.1) of the normal lives of organisms alter contemporary evolutionary psychologies? Most significantly, it would disconnect how evolutionary science approaches the study of behaviour from any constraint by ideas about how natural selection operates (e.g. the need to calculate ‘inclusive fitness’; or to speculate about a prehistoric ‘environment of evolutionary adaptedness’; Buss, 2009 ; Tooby & Cosmides, 2016 ). Because, as Walsh ( 2015 , Ch.2.1) says, ‘given the normal activities of organisms, nothing needs to be added [to our theoretical framework] to get populations to change in the ways that Darwin describes as natural selection.’ Which represents a complete reversal of those tenets of evolutionary psychology that produce statements like these (italics mine):

Like vision and language, our emotions and cognitive faculties are complex, useful, and non-randomly organized, which means that they must be a product of the only physical process capable of generating complex, useful, non-random organization, namely, natural selection (Pinker, 2005 , p. xiv).

Because mental phenomena are the expression of complex functional organization in biological systems, and complex organic functionality is the downstream consequence of natural selection , then it must be the case that the sciences of the mind and brain are adaptationist sciences, and psychological mechanisms are computational adaptations (Tooby & Cosmides, 2016 , p. 11).

[The brain’s] programs were designed not by an engineer, but by natural selection, a causal process that retains and discards design features based on how well they solved adaptive problems in past environments (Tooby & Cosmides, 2016 , p. 19).

If natural selection is not an ‘upstream’ causal process which produces psychological phenomena, but a ‘downstream consequence’ of the normal activities of organisms—only some of which have adaptive consequences (cf. Chauncey Wright, above)—then our theoretical attention must switch from claims about natural selection, to the need adequately to conceptualise how agency manifests itself in the natural world. Perhaps we should not be surprised, therefore, that when we examine how Darwin himself presented his studies of various creatures’ ‘habits’ or behaviour—human group-processes and facial expressions; sexual displays in animals; worms’ intelligence and the motility of the ova of Flustra ; mutual aid among social animals and the problem-solving movements of plant growth—we find these all reflect a single, coherent vision. According to Darwin (e.g. 1859a, p. 61), any studied habitat is maintained by the ‘infinitely complex’ web of actions and reactions linking the habits of the focal organism ‘to other organic beings and to external nature.’ It is this vision of the interdependencies created and maintained by agency which underpins how Darwin construed what he called ‘the struggle for life,’ and, as a consequence, how he understood natural selection (Bradley, 2020 ).

The idea that natural selection is the causal force or mechanism which produces evolutionary adaptations and originates new species remains for many a scientific truism, thanks to the continuing appeal of ‘genes-eye’ MS accounts of evolution. This essay rejects a corresponding truism in Darwin scholarship, which holds that the main aim of Origin ’s (Darwin, 1859a , p. 459) ‘one long argument’ is to prove natural selection the causal mechanism or vera causa responsible for the evolution of adaptations and new species. 14 Specifically, I show how modern historiographic constructions of Darwin’s supposed authorial ‘intention,’ ‘desperation,’ or ‘commitment’ to prove natural selection a vera causa in Origin are built on an unnecessarily selective sample from what Darwin himself wrote about verae causae , typically highlighting just one remark, comprising the first third of Darwin’s ( 1863b ) brief postscript to a letter to George Bentham.

My starting-point was different. I began by examining the context for all Darwin’s known uses of the term ‘ vera causa .’ From this beginning I have argued that, provided one grounds one’s views of Origin ’s arguments upon: how Darwin himself used the term vera causa (as requiring first-hand observational evidence); how this usage conforms to the commonest meaning of the term among Victorian philosophers of science; how Origin itself sets up, and repeatedly restates, the logical dependence of natural selection on inheritance, variation and the struggle for life; how, responding to criticism early in 1860, Darwin disputed the need to prove natural selection the true cause of adaptive change and evolution; and how that dispute led him to revise later editions of the book—then one must conclude that, according to Darwin, natural selection is an effect of other causes, not a cause in its own right.

One advantage of recognising that Origin does not comprise just one argument—aimed at proving natural selection the true cause of adaptation—is to re-focus historiographic and scientific attention on all the other arguments that the book makes. Several of these arguments have become central to evolutionary science over the last twenty years, though often without any awareness by modern scientists of antecedent arguments in Origin . 15 These include the leading role played by organisms’ agency in the genesis of adaptations (cf. ‘transitional habits’ aka the ‘Baldwin effect,’ ‘genetic accommodation,’ and ‘niche construction’: Darwin, 1876 , pp. 138–143; Gould, 2002 , pp. 125–127; Noble & Noble, 2017 ; Noble, 2021 ; Odling-Smee et al., 2003 ; Walsh, 2015 ); the importance of plasticity of structure in the evolution of new adaptations (Darwin, 1876 , pp. 62, 106, 438; West-Eberhard, 2003 ; 2008 ); the direct effect of external conditions (Darwin, 1876 , p. 67; Gilbert & Epel, 2015 , pp. 435ff); the recognition that ‘inheritance’ includes ‘two distinct processes’—the transmission of heritable characters from parent to offspring and their development (Darwin, 1876 , pp. 114–15, 119–122; 1874 , p. 227; Walsh, 2010 ); and multi-level selection (Darwin, 1876 , pp. 67–68; Wilson & Wilson, 2007 ).

As soon as contemporary scientists accept that, as per Darwin’s argument in Origin , natural selection does not cause, but results from the ordinary activities of organisms, contemporary evolutionary theorists must address a new foundational challenge: the need to construct a viable, evidence-based picture of the natural world as what I have called a ‘theatre of agency’ (Bradley, 2020 ). Only when they have such a picture will scientists be in a position to work up an intelligible account of natural selection. The pioneering instance of such a working-up constitutes a theme central to Darwin’s many publications.

Acknowledgements

With thanks to Michael Ruse for a stimulating email correspondence on this topic, and to Jane Selby for help and discussion throughout.

Open Access funding enabled and organized by CAUL and its Member Institutions.

Declarations

I have no conflicts of interest. Research was self-funded.

1 E.g. the third edition of Origin (Darwin, 1861a , p. 88, my italics) adjusted the above quotation to begin: ‘ It may metaphorically be said that natural selection is daily and hourly scrutinising, throughout the world …’.

2 Physicists still debate what causes the phenomena the law of gravity describes. At least fourteen theories of gravity are currently in play.

3 NB Darwin here denies that the undulatory theory of light identifies a ‘vera causa’ on the grounds that ethereal undulations had never been observed. In this he equates verae causae with demonstration by positive observational evidence (as discussed later in this section). Conversely, Darwin also implies that causal hypotheses (e.g. ‘interference patterns result from the wave-form of light’) do not have to equate to verae causae . Hence the need for my examination of the way the argument in Origin is presented to prove my case that he cast natural selection as a consequence of other causes (see Sect.  5.2 below).

4 Note that controversy over the causal status of natural selection was still live in 1870: ‘Strictly speaking, Natural Selection is not a cause at all, but is the mode of operation of a certain quite limited class of causes’: Chauncey Wright, ‘Review [of Contributions to the Theory of Natural Selection. A Series of Essays by Alfred Russell Wallace],’ North American Review111, (1870), p. 293.

5 It is notable that this postscript is never quoted in full by the authors I have cited. All stop before the words, ‘When we descend to details …’ Even I have not quoted its two final sentences, viz. ‘The latter case seems to me hardly more difficult to understand precisely & in detail than the former case of supposed change. Bronn may ask in vain the old creationist school & the new school why one mouse has longer ears than another mouse—& one plant more pointed leaves than another plant.'

6 Hull ( 1973 , pp. 7–8; 2003 , pp. 185–188) argues that Darwin was mistaken in this belief, at least where Mill was concerned.

7 Albeit in its second paragraph, which historiographers seldom if ever quote or discuss—as noted previously.

8 A contributing factor here may have been Darwin’s growing sense that his Glen Roy ( 1839 ) paper had been a ‘gigantic blunder.’ According to Barrett ( 1973 , p. 24) Darwin came to the conclusion that it was ‘metaphysics’ that ‘had tripped him.’ Hence his determination that this ‘didn’t happen again.’.

9 This eruption gave birth to Sabrina Island.

10 This difficulty was even more acute than with Newton’s law of gravity. Yes, gravity was indeed invisible, and had an unknown causal mechanism. But its putative effects can be observed and measured here and now: pendulums swinging; apples accelerating towards earth; tides rising and falling; planets orbiting. Natural selection could neither be observed in itself. Nor could the immensely slow production of its putative effects be observed in the span of any one human lifetime.

11 Or, indeed, arguably from its title , which equates, ‘the origin of species by means of natural selection,’ with ‘the preservation of favoured races in the struggle for life.’

12 A causal mechanism for inheritance is tentatively presented in the last chapter of Variation under Domestication (Darwin, 1868 ), titled: ‘Provisional hypothesis of pangenesis,’ though this hypothesis was never incorporated or mentioned in later editions of Origin.

13 This process was supposedly only discovered after Darwin’s death, by James Mark Baldwin in 1896 (similar formulations were advanced at about the same time by Conway Lloyd Morgan and others). As a result it is now known as ‘the Baldwin effect’ (after Simpson, 1953 ). However, Baldwin ( 1909 , p. 11) explicitly denied that he had discovered this process, which he called ‘organic selection,’ because, he said, Origin had clearly described it in 1859, under the heading ‘transitional habits.’

14 The very idea that this claim of Darwin’s ( 1859a , p. 459)—that Origin comprised ‘one long argument’—espoused a sober epistemological commitment, arguably misrecognises its rhetorical, not to say wishful, status. In her commentary, Beer’s ( 2000 , pp. 42–43) reminds us how Origin ’s ‘argument’ does not proceed according to an epistemological schema, but rather by laboriously accumulating ‘an unruly superfluity of material’ through ‘a strange intermingling of acquisition, concretion, analogy and prophecy.’ Hence, it is ‘only gradually and retrospectively,’ she says, that ‘the force of the [‘one long’] argument emerge[s] from the profusion of example.’.

15 There are several exceptions to this claim, most notably for me; Mary Jane West-Eberhard ( 2003 , 2008 ).

The origiinal online version of this article was revised: Missing Open Access funding information has been added in the Funding Note.

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The "Natural Order" of Morpheme Acquisition: A Historical Survey and Discussion of Three Putative Determinants

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Akira Murakami

We revisit morpheme studies to evaluate the long-standing claim for a universal order of acquisition. We investigate the L2 acquisition order of six English grammatical morphemes by learners from seven L1 groups across five proficiency levels. Data are drawn from approximately 10,000 written exam scripts from the Cambridge Learner Corpus. The study establishes clear L1 influence on the absolute accuracy of morphemes and their acquisition order, therefore challenging the widely held view that there is a universal order of acquisition of L2 morphemes. Moreover, we find that L1 influence is morpheme specific, with morphemes encoding language-specific concepts most vulnerable to L1 influence.

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This study sought to determine the sequence of L2 morpheme presentation, as well as to determine whether or not the sequence of morpheme presentations correspond with the recognized natural order of morpheme acquisition in English Language Teaching course books utilized with young adult learners at a public sector vocational education institution in Thailand. Qualitative analysis was employed in the scrutinizing of twelve beginner and elementary level ESL and EFL learners course books that have been utilized as the primary teaching material for over a decade by the general education department of the institute. This examination revealed that the morpheme presentation sequence within the selected ELT course books was not analogous with the conclusions in the supporting literature. The findings further indicated that the widely accepted viewpoint of natural order morpheme acquisition was likewise not substantially reflected within the analyzed texts. Albeit, earlier studies have found that an unnatural sequence of morpheme presentation in EFL course books may hamper communicative competence in English, further study is required to establish if this may be a contributing factor for the overall low English proficiency of adult L2 learners in Thailand.

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Ever since Dulay and Burt (1974), it is believed that children learning English as a Second Language, irrespective of their learning context, follow a fixed order of acquisition of English grammatical morphemes. However, works like Luk and Shirai (2009) have shown that the effects of L1 in language learning is very strong and despite the numerous claims for the 'natural' order of L2 acquisition of English grammatical morphemes, they are no exception to L1 influence. Keeping in mind the contentions of the previous studies done in the area of acquisition order of English grammatical morphemes, this paper examines the accuracy order of five English grammatical morphemes (plural-s, irregular past, regular past and articles) of L1 Mizo learners of English as a Second Language. A formula for Target Like Use (TLU) score taken from Pica (1983) is used for analysis of twenty-seven scripts derived from twenty-seven participants. The result of the study indicates that there is a possible L1 influence in the L2 acquisition order of English grammatical morphemes; however, a larger cross-linguistic based study can be done for generalization of the results.

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The present study examined the proposal that the presence of a similar morpheme in the learner's first and second languages (L2) facilitates morphological development in the L2. Advanced Russian and Japanese speakers of English as a second language performed a self-paced reading task in which they read English sentences word by word for comprehension. Russian participants showed a reliable sensitivity to plural errors, but Japanese participants did not. The findings supported the morphological congruency hypothesis. A theoretical proposal is put forward to explain how morphological con-gruency affects L2 morpheme acquisition. The findings and the proposal are relevant to the discussion of the critical period hypothesis, ultimate attainment in L2, and the characterization of L2 competence of steady-state adult L2 learners. The acquisition of grammatical morphemes by second-language (L2) learners has received a great deal of attention in SLA research, both as a research topic in its own right and as a means to investigate other SLA issues such as the aspect

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Understanding the Core Principles of Natural Law

This essay about natural law explores its fundamental principles, historical development, and relevance in modern ethical and legal discussions. Originating from Greek and Roman philosophy, natural law asserts that inherent principles govern morality and legality, discernible through human reason. Influential thinkers like Aristotle, Cicero, and Aquinas contributed to its evolution, integrating it with Christian doctrine. Key tenets include universality, objective morality, rational discoverability, and promotion of the common good. Despite criticisms, natural law remains significant in debates on human rights and legal interpretations, emphasizing that true justice aligns with an intrinsic moral order.

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Greek thinkers so as for example Aristotle and Plato set aside suggestion foundation, that command inalienable universe. Aristotle put a concept ‘justice, that outstrips the laws done man nature’, while Plato theory unreplacement brings up or ideals offered a moral higher order.

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Department of Cognitive Science

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Hongru Zhu (PhD ’23) published a paper with Prof. Daniel Kersten (UMN) on Natural scenes reveal diverse representations of 2D and 3D body pose in the human brain  in Proceedings of the National Academy of Sciences based on research he started as a student in our program.

https://www.pnas.org/doi/10.1073/pnas.2317707121

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Research on the detection method of the threshing rate of corn ears based on machine vision, share and cite.

Li, X.; Xu, S.; Zhang, W.; Wang, J.; Li, Y.; Peng, B.; Sun, R. Research on the Detection Method of the Threshing Rate of Corn Ears Based on Machine Vision. Agriculture 2024 , 14 , 1037. https://doi.org/10.3390/agriculture14071037

Li X, Xu S, Zhang W, Wang J, Li Y, Peng B, Sun R. Research on the Detection Method of the Threshing Rate of Corn Ears Based on Machine Vision. Agriculture . 2024; 14(7):1037. https://doi.org/10.3390/agriculture14071037

Li, Xinping, Shendi Xu, Wantong Zhang, Junyi Wang, Yanan Li, Bin Peng, and Ruizhe Sun. 2024. "Research on the Detection Method of the Threshing Rate of Corn Ears Based on Machine Vision" Agriculture 14, no. 7: 1037. https://doi.org/10.3390/agriculture14071037

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