Evolution in Darwin's finches: a review of a study on Isla Daphne Major in the Galápagos archipelago

This paper reviews research pertaining to the problem of speciation of the finches on the Galápagos archipelago carried out by assistants, several colleagues, Peter Grant and myself. I give a brief history of the radiation, examine the process of divergence by natural selection over time, and then consider the nature of the reproductive barrier to gene flow between closely related species. Fluctuating climatic conditions have produced a continuously changing ecological landscape and altered feeding conditions for the finches over the last 30 years. Finch populations tracked these changes by natural selection and evolutionary responses to the main events. At each event significant morphological change occurred from one generation to the next generation. As a consequence of these accumulated changes, the mean bill shape and body size of the Geospiza fortis and G. scandens populations differed markedly from 1973 to 2002. Song, a learned culturally transmitted trait, acted as a barrier to reproduction between these species. Rare incidences of misimprinting on song led to hybridization and introgression. Low levels of gene flow from one species to another increased genetic variation on which selection acted. Although the major driving force of diversification was ecological change, the process of diversification involved a subtle interplay between ecology, genetic evolution and learned culturally transmitted traits. An important message for conservation is that neither the environment nor species are fixed entities, therefore a wise strategy for conserving endangered species should keep them capable of further change.     

Foreseeing fates: a commentary on Manton (1928) ‘On the embryology of a mysid crustacean,Hemimysis lamornae’

Sidnie Manton became best known for her work on arthropod locomotion, and for proposing radical views on the evolution of arthropods that were accepted for a generation. However, her early training was as an embryologist, and the work that she carried out at the beginning of her career still stands as one of the major twentieth century contributions to the study of crustacean embryology. Here, I review her first major paper, largely completed while she was a graduate student, describing embryonic development in Hemimysis lamornae, a small shrimp-like animal found in the seas around the UK. The clarity of her writing and the quality of her figures set a standard that laid the basis for subsequent work, and although not all of her conclusions have stood the test of time, they remain a standard reference for work today. This commentary was written to celebrate the 350th anniversary of the journal Philosophical Transactions of the Royal Society.     

The prevention and eradication of smallpox: a commentary on Sloane (1755) ‘An account of inoculation’

Sir Hans Sloane''s account of inoculation as a means to protect against smallpox followed several earlier articles published in Philosophical Transactions on this procedure. Inoculation (also called ‘variolation’) involved the introduction of small amounts of infectious material from smallpox vesicles into the skin of healthy subjects, with the goal of inducing mild symptoms that would result in protection against the more severe naturally acquired disease. It began to be practised in England in 1721 thanks to the efforts of Lady Mary Wortley Montagu who influenced Sloane to promote its use, including the inoculation of the royal family''s children. When Edward Jenner''s inoculation with the cow pox (‘vaccination’) followed 75 years later as a safer yet equally effective procedure, the scene was set for the eventual control of smallpox epidemics culminating in the worldwide eradication of smallpox in 1977, officially proclaimed by WHO in 1980. Here, we discuss the significance of variolation and vaccination with respect to scientific, public health and ethical controversies concerning these ‘weapons of mass protection’. This commentary was written to celebrate the 350th anniversary of the journal Philosophical Transactions of the Royal Society.     

Deciphering death: a commentary on Gompertz (1825) ‘On the nature of the function expressive of the law of human mortality,and on a new mode of determining the value of life contingencies’

In 1825, the actuary Benjamin Gompertz read a paper, ‘On the nature of the function expressive of the law of human mortality, and on a new mode of determining the value of life contingencies’, to the Royal Society in which he showed that over much of the adult human lifespan, age-specific mortality rates increased in an exponential manner. Gompertz''s work played an important role in shaping the emerging statistical science that underpins the pricing of life insurance and annuities. Latterly, as the subject of ageing itself became the focus of scientific study, the Gompertz model provided a powerful stimulus to examine the patterns of death across the life course not only in humans but also in a wide range of other organisms. The idea that the Gompertz model might constitute a fundamental ‘law of mortality’ has given way to the recognition that other patterns exist, not only across the species range but also in advanced old age. Nevertheless, Gompertz''s way of representing the function expressive of the pattern of much of adult mortality retains considerable relevance for studying the factors that influence the intrinsic biology of ageing. This commentary was written to celebrate the 350th anniversary of the journal Philosophical Transactions of the Royal Society.     

Forging patterns and making waves from biology to geology: a commentary on Turing (1952) ‘The chemical basis of morphogenesis’

Alan Turing was neither a biologist nor a chemist, and yet the paper he published in 1952, ‘The chemical basis of morphogenesis’, on the spontaneous formation of patterns in systems undergoing reaction and diffusion of their ingredients has had a substantial impact on both fields, as well as in other areas as disparate as geomorphology and criminology. Motivated by the question of how a spherical embryo becomes a decidedly non-spherical organism such as a human being, Turing devised a mathematical model that explained how random fluctuations can drive the emergence of pattern and structure from initial uniformity. The spontaneous appearance of pattern and form in a system far away from its equilibrium state occurs in many types of natural process, and in some artificial ones too. It is often driven by very general mechanisms, of which Turing''s model supplies one of the most versatile. For that reason, these patterns show striking similarities in systems that seem superficially to share nothing in common, such as the stripes of sand ripples and of pigmentation on a zebra skin. New examples of ‘Turing patterns'' in biology and beyond are still being discovered today. This commentary was written to celebrate the 350th anniversary of the journal Philosophical Transactions of the Royal Society.     

Focusing by shape change in the lens of the eye: a commentary on Young (1801) ‘On the mechanism of the eye’

In his Bakerian Lecture paper of 1801, Thomas Young provided the best account up to that time of the eye''s optical system, including refraction by the cornea and the surfaces of the lens. He built a device, an optometer, for determining the eye''s state of focus, making it possible to prescribe appropriate correction lenses. His main contribution, however, was to show that accommodation, the eye''s focusing mechanism, was not the result of changes to the curvature of the cornea, nor to the length of the eye, but was due entirely to changes in the shape of the lens, which he described with impressive accuracy. He was wrong, however, in believing that the reason the lens bulges when focusing on near objects was because it behaved as a contracting muscle. Half a century later, Helmholtz showed that the lens bulges not by its own contraction, but when it is relaxed as a result of contraction of newly discovered circular muscles in the ciliary body. This commentary was written to celebrate the 350th anniversary of the journal Philosophical Transactions of the Royal Society.     

How mathematical epidemiology became a field of biology: a commentary on Anderson and May (1981) ‘The population dynamics of microparasites and their invertebrate hosts’

We discuss the context, content and importance of the paper ‘The population dynamics of microparasites and their invertebrate hosts’, by R. M. Anderson and R. M. May, published in the Philosophical Transactions of the Royal Society as a stand-alone issue in 1981. We do this from the broader perspective of the study of infectious disease dynamics, rather than the specific perspective of the dynamics of insect pathogens. We argue that their 1981 paper fits seamlessly in the systematic study of infectious disease dynamics that was initiated by the authors in 1978, combining effective use of simple mathematical models, firmly rooted in biology, with observable or empirically measurable ingredients and quantities, and promoting extensive capacity building. This systematic approach, taking ecology and biology rather than applied mathematics as the motivation for advance, proved essential for the maturation of the field, and culminated in their landmark textbook of 1991. This commentary was written to celebrate the 350th anniversary of the journal Philosophical Transactions of the Royal Society.     

Extraordinary fossils reveal the nature of Cambrian life: a commentary on Whittington (1975) ‘The enigmatic animal Opabinia regalis,Middle Cambrian,Burgess Shale,British Columbia’

Harry Whittington''s 1975 monograph on Opabinia was the first to highlight how some of the Burgess Shale animals differ markedly from those that populate today''s oceans. Categorized by Stephen J. Gould as a ‘weird wonder’ (Wonderful life, 1989) Opabinia, together with other unusual Burgess Shale fossils, stimulated ongoing debates about the early evolution of the major animal groups and the nature of the Cambrian explosion. The subsequent discovery of a number of other exceptionally preserved fossil faunas of Cambrian and early Ordovician age has significantly augmented the information available on this critical interval in the history of life. Although Opabinia initially defied assignment to any group of modern animals, it is now interpreted as lying below anomalocaridids on the stem leading to the living arthropods. This commentary was written to celebrate the 350th anniversary of the journal Philosophical Transactions of the Royal Society.     

A model model: a commentary on DiFrancesco and Noble (1985) ‘A model of cardiac electrical activity incorporating ionic pumps and concentration changes’

This paper summarizes the advances made by the DiFrancesco and Noble (DFN) model of cardiac cellular electrophysiology, which was published in Philosophical Transactions B in 1985. This model was developed at a time when the introduction of new techniques and provision of experimental data had resulted in an explosion of knowledge about the cellular and biophysical properties of the heart. It advanced the cardiac modelling field from a period when computer models considered only the voltage-dependent channels in the surface membrane. In particular, it included a consideration of changes of both intra- and extracellular ionic concentrations. In this paper, we summarize the most important contributions of the DiFrancesco and Noble paper. We also describe how computer modelling has developed subsequently with the extension from the single cell to the whole heart as well as its use in understanding disease and predicting the effects of pharmaceutical interventions. This commentary was written to celebrate the 350th anniversary of the journal Philosophical Transactions of the Royal Society.     

The new nativism: a commentary on Gary Marcus’s The birth of the mind

Gary Marcus (2004) has written a very interesting book about mental development from a nativist perspective. For the general readership at which the book is largely aimed, it will be interesting because of its many informative examples of the development of cognitive structures and because of its illuminating explanations of ways in which genes can contribute to these developmental processes. However, the book is also interesting from a theoretical point of view. Marcus tries to make nativism compatible with the central arguments that anti-nativists use to attack nativism and with many recent discoveries about genetic activity and brain development. In so doing, he reconfigures the nativist position to a considerable extent. Marcus’s theory is certainly more sophisticated than any version of nativism on the market. However, in our view Marcus ends up reconfiguring the nativist position out of existence. While many of the points that Marcus makes are both interesting and correct, we see no compelling reason to classify his considered position with traditional nativism rather than anti-nativism. More generally, we that think his book points to a general moral: the opposition between nativism and anti-nativism does not help us to understand psycho-developmental issues, and should therefore be abandoned.     

Effects of laboratory colonization on Bactrocera dorsalis (Diptera,Tephritidae) mating behaviour: ‘what a difference a year makes’

Laboratory-reared insects are widely known to have significantly reduced genetic diversity in comparison to wild populations; however, subtle behavioural changes between laboratory-adapted and wild or ‘wildish’ (i.e., within one or very few generations of field collected material) populations are less well understood. Quantifying alterations in behaviour, particularly sexual, in laboratory-adapted insects is important for mass-reared insects for use in pest management strategies, especially those that have a sterile insect technique component. We report subtle changes in sexual behaviour between ‘wildish’ Bactrocera dorsalis flies (F1 and F2) from central and southern Thailand and the same colonies 12 months later when at six generations from wild. Mating compatibility tests were undertaken under standardised semi-natural conditions, with number of homo/heterotypic couples and mating location in field cages analysed via compatibility indices. Central and southern populations of Bactrocera dorsalis displayed positive assortative mating in the 2010 trials but mated randomly in the 2011 trials. ‘Wildish’ southern Thailand males mated significantly earlier than central Thailand males in 2010; this difference was considerably reduced in 2011, yet homotypic couples from southern Thailand still formed significantly earlier than all other couple combinations. There was no significant difference in couple location in 2010; however, couple location significantly differed among pair types in 2011 with those involving southern Thailand females occurring significantly more often on the tree relative to those with central Thailand females. Relative participation also changed with time, with more southern Thailand females forming couples relative to central Thailand females in 2010; this difference was considerably decreased by 2011. These results reveal how subtle changes in sexual behaviour, as driven by laboratory rearing conditions, may significantly influence mating behaviour between laboratory-adapted and recently colonised tephritid fruit flies over a relatively short period of time.     

From plant extract to molecular panacea: a commentary on Stone (1763) ‘An account of the success of the bark of the willow in the cure of the agues’

The application of aspirin-like drugs in modern medicine is very broad, encompassing the treatment of inflammation, pain and a variety of cardiovascular conditions. Although anecdotal accounts of willow bark extract as an anti-inflammatory drug have occurred since written records began (for example by Hippocrates), the first convincing demonstration of a potent anti-pyretic effect of willow bark containing salicylates was made by the English cleric Edward Stone in the late eighteenth century. Here, we discuss the route to optimizing and understanding the mechanism of action of anti-inflammatory drugs that have their origins in Stone''s seminal study, ‘An account of the success of the bark of the willow in the cure of agues’. This commentary was written to celebrate the 350th anniversary of the journal Philosophical Transactions of the Royal Society.     

Spawning aggregations of squid (Sepioteuthis australis) populations: a continuum of ‘microcohorts’

The aim of this study was to determine how size, age, somatic and reproductive condition, abundance and egg production of southern calamary spawning aggregations changed during the spawning season in each of 2 years. During the spawning period in at least one of the years there was a decline as much as 20% in average size, 50% in somatic condition, 28–34% in size-at-age, 26–29% in reproductive status, as well as abundance and reproductive output of the stock declining during the spawning season. However, this change was not a function of the population becoming reproductively exhausted, as the aggregation was composed of different individuals with different biological characteristics. In each month the average age of individuals was ca. 6 mo, indicating that squid that had hatched at different times had entered the spawning aggregations, suggesting that the aggregation was made-up of a succession of microcohorts. Currently, management of many squid populations assumes that there is a single cohort in the aggregation. Therefore, estimating stock biomass at the start of the spawning season cannot be used as the population is constantly changing as micro-cohorts move into the aggregation. An instantaneous estimate of the spawning biomass, independent of fishing activity may be obtained by quantifying the density of deposited eggs. The strategy of individuals with a diversity of life history characteristics coming together in a single spawning aggregation may ensure the phenotypic and genetic diversity required to guarantee successful recruitment of this short-lived species. Therefore, temporally structured protection from harvest throughout the spawning season will ensure maintenance of this population diversity.     

The early evolution of land plants,from fossils to genomics: a commentary on Lang (1937) ‘On the plant-remains from the Downtonian of England and Wales'

During the 1920s, the botanist W. H. Lang set out to collect and investigate some very unpromising fossils of uncertain affinity, which predated the known geological record of life on land. His discoveries led to a landmark publication in 1937, ‘On the plant-remains from the Downtonian of England and Wales’, in which he revealed a diversity of small fossil organisms of great simplicity that shed light on the nature of the earliest known land plants. These and subsequent discoveries have taken on new relevance as botanists seek to understand the plant genome and the early evolution of fundamental organ systems. Also, our developing knowledge of the composition of early land-based ecosystems and the interactions among their various components is contributing to our understanding of how life on land affects key Earth Systems (e.g. carbon cycle). The emerging paradigm is one of early life on land dominated by microbes, small bryophyte-like organisms and lichens. Collectively called cryptogamic covers, these are comparable with those that dominate certain ecosystems today. This commentary was written to celebrate the 350th anniversary of the journal Philosophical Transactions of the Royal Society.