1 July 1858 and the 1844 essay: what Lyell and Hooker decided; and what Darwin did not want and did not know

Two ‘papers’ of Darwin's were read at the famous 1 July 1858 meeting of the Linnean Society: an excerpt from his 1844 essay and a summary of his theory, enclosed in an 1857 letter to Asa Gray. Quite apart from not selecting the essay excerpt, Darwin's letters appear to indicate that he definitely did not want, and hence did not expect, an excerpt from his 1844 essay to be included (and that he did not learn of its inclusion until some 2 weeks after the meeting). As a result, we refine Darwin's role in ‘the delicate arrangement’, as well as the basis for Hooker's and Lyell's. In particular, why did they choose an essay excerpt to be presented contrary to Darwin's wishes? In direct opposition to the popular view, the essay excerpt was the afterthought, the last‐minute add‐on, not the enclosure to the Gray letter.     

Darwin and barnacles

In this essay, I discuss the origin of Charles Darwin's interest in cirripedes (barnacles). Indeed, he worked intensively on cirripedes during the years in which he was developing the theory that eventually led to the publication of The Origin of Species. In the light of our present knowledge, I present Darwin's achievements in the morphology, systematics and biology of these small marine invertebrates, and also his mistakes. I suggest that the word that sheds the most light here is homology, and that his mistakes were due to following Richard Owen's method of determining homologies by reference to an ideal archetype. I discuss the ways in which his studies on cirripedes influenced the writing of The Origin.     

Darwin and palaeontology: a re-evaluation of his interpretation of the fossil record

Charles Darwin's empirical research in palaeontology, especially on fossil invertebrates, has been relatively neglected as a source of insight into his thinking, other than to note that he viewed the fossil record as very incomplete. During the Beagle voyage, Darwin gained extensive experience with a wide diversity of fossil taxa, and he thought deeply about the nature of the fossil record. That record was, for him, a major source of evidence for large-scale transmutation, but much less so for natural selection or single lineages. Darwin's interpretation of the fossil record has been criticised for its focus on incompleteness, but the record as he knew it was extremely incomplete. He was compelled to address this in arguing for descent with modification, which was likely his primary goal. Darwin's gradualism has been both misrepresented and exaggerated, and has distracted us from the importance of the fossil record in his thinking, which should be viewed in the context of the multiple, sometimes competing demands of the multifaceted argument he presented in the Origin of Species.     

Sexual selection: Another Darwinian process

Why was sexual selection so important to Darwin? And why was it de-emphasized by almost all of Darwin's followers until the second half of the 20th century? These two questions shed light on the complexity of the scientific tradition named “Darwinism”. Darwin's interest in sexual selection was almost as old as his discovery of the principle of natural selection. From the beginning, sexual selection was just another “natural means of selection”, although different from standard “natural selection” in its mechanism. But it took Darwin 30 years to fully develop his theory, from the early notebooks to the 1871 book The Descent of Man, and Selection in Relation to Sex. Although there is a remarkable continuity in his basic ideas about sexual selection, he emphasized increasingly the idea that sexual selection could oppose the action of natural selection and be non adaptive. In time, he also gave more weight to mate choice (especially female choice), giving explicit arguments in favor of psychological notions such as “choice” and “aesthetic sense”. But he also argued that there was no strict demarcation line between natural and sexual selection, a major difficulty of the theory from the beginning. Female choice was the main reason why Alfred Russel Wallace, the co-discoverer of the principle of natural selection, engaged in a major controversy with Darwin about sexual selection. Wallace was suspicious about sexual selection in general, trying to minimize it by all sorts of arguments. And he denied entirely the existence of female choice, because he thought that it was both unnecessary and an anthropomorphic notion. This had something to do with his spiritualist convictions, but also with his conception of natural selection as a sufficient principle for the evolutionary explanation of all biological phenomena (except for the origin of mind). This is why Wallace proposed to redefine Darwinism in a way that excluded Darwin's principle of sexual selection. The main result of the Darwin–Wallace controversy was that most Darwinian biologists avoided the subject of sexual selection until at least the 1950 s, Ronald Fisher being a major exception. This controversy still deserves attention from modern evolutionary biologists, because the modern approach inherits from both Darwin and Wallace. The modern approach tends to present sexual selection as a special aspect of the theory of natural selection, although it also recognizes the big difficulties resulting from the inevitable interaction between these two natural processes of selection. And contra Wallace, it considers mate choice as a major process that deserves a proper evolutionary treatment. The paper's conclusion explains why sexual selection can be taken as a test case for a proper assessment of “Darwinism” as a scientific tradition. Darwin's and Wallace's attitudes towards sexual selection reveal two different interpretations of the principle of natural selection: Wallace's had an environmentalist conception of natural selection, whereas Darwin was primarily sensitive to the element of competition involved in the intimate mechanism of any natural process of selection. Sexual selection, which can lack adaptive significance, reveals this exemplarily.     

Haeckel,un darwinien allemand ?

German biologist Ernst Haeckel (1834–1919) is often considered the most renowned Darwinian in his country since, as early as 1862, he declared that he accepted the conclusions Darwin had reached three years before in On the Origin of Species, and afterwards, he continuously proclaimed himself a supporter of the English naturalist and championed the evolutionary theory. Nevertheless, if we examine carefully his books, in particular his General Morphology (1866), we can see that he carries on a tradition very far from Darwin's thoughts. In spite of his acceptance of the idea of natural selection, that he establishes as an argument for materialism, he adopts, indeed, a conception of evolution that is, in some respects, rather close to Lamarck's views. He is, thus, a good example of the ambiguities of the reception of Darwinism in Germany in the second part of the 19th century. To cite this article: S. Schmitt, C. R. Biologies 332 (2009).     

Darwin and sexual selection: One hundred years of misunderstanding

Darwin's book on the Descent of Man and Selection in Relation to Sex (1871) is often viewed as the continuation of TheOrigin of Species published 12 years earlier (1859), both because of the implicit parallelism between natural selection and sexual selection, and because Darwin himself presents the book as developing a subject (man) which he intentionally omitted in the Origin. But the Descent can also be viewed as the continuation of his book on Variation published three years earlier (1868). Firstly because Darwin's hypothesis of pangenesis links the selection process to the origin of variation through use and disuse, an idea underlying his speculations on the origin of moral sense in humans. Second because like the action of the horticulturist on his domestic crops, sexual selection exerted by one sex on the other sex can develop fancy traits that are not easily accounted for by their utility to the selected organism itself, such as artistic taste, pride, courage, and the morphological differences between human populations. These traits are difficult to reconcile with pangenesis. They add up to other contradictions of the book possibly resulting from Darwin's erroneous inference about the mechanism of inheritance, like those on the determination of sex-ratio, or the confusion between individual adaptation and the advantage to the species. These inconsistencies inaugurate a weakening of the Darwinian message, which will last 50 years after his death. They contributed to the neglect of sexual selection for a century. Darwin however maintained a logical distinction between evolutionary mechanisms and hereditary mechanisms, and an epistemological distinction between evolutionary theory and Pangenesis hypothesis. In the modern context of Mendelian genetics, Darwin's sexual selection retrospectively appears as luminous an idea in its pure principle as natural selection, even though the mechanisms governing the evolution of sexual choice in animals remain largely unresolved.     

Epistemology, necessity, and evolution: a critical review of Michael Ruse��s Philosophy After Darwin

Michael Ruse??s new anthology Philosophy After Darwin provides great history and background in the major impacts Darwinism has had on philosophy, especially in ethics and epistemology. This review focuses on epistemology understood through the lens of evolution by natural selection. I focus on one of Ruse??s own articles in the collection, which responds to two classic articles by Konrad Lorenz and David Hull on the two major forms of evolutionary epistemology. I side with Ruse against Lorenz??s account of the necessity we think our principles of reasoning have, though I disagree with Ruse??s particular example. I also argue that Ruse??s alternative explanation is lacking. Against Hull, I side with Ruse in his doubts that a sociobiological approach to science will prove fruitful, though I point out that it has certain advantages other approaches do not have. Although I side with Ruse on the issue, I conclude that the two views do not really come into direct conflict and so one needs not reject either. Finally, I discuss Ruse??s positive view and raise questions for his conception of evolutionary epistemology. I conclude that his arguments are insufficient to overcome opposing views and his view has at least as many unintuitive conclusions as the alternatives.     

Darwin's puzzling Expression

Charles Darwin's The Expression of the Emotions in Man and Animals (1872) is a very different kind of work from On the Origin of Species (1859). This “otherness” is most extreme in the character of the explanations that Darwin offers in the Expression. Far from promoting his theory of natural selection, the Expression barely mentions that theory, instead drawing on explanatory principles which recall less Darwinian than Lamarckian and structuralist biological theorizing. Over the years, historians have offered a range of solutions to the puzzle of why the Expression is so “non-Darwinian”. Close examination shows that none of these meets the case. However, recent research on Darwin's lifelong engagement with the controversies in his day over the unity of the human races makes possible a promising new solution. For Darwin, emotional expression served the cause of defending human unity precisely to the extent that natural selection theory did not apply.     

Darwin teleologist? Design in the Orchids

Focusing on the Orchids, this article aims at disentangling the concepts of teleology, design and natural theology. It refers to several contemporary critics of Darwin (Kölliker, Argyll, Royer, Candolle, Delpino) to challenge Huxley's interpretation that Darwin's system was “a deathblow” to teleology. The Orchids seem rather to be a “flank-movement” (Gray): it departs from the Romantic theories of transmutation and the “imaginary examples” of the Origin; it focuses on empirical data and on teleological structures. Although Darwin refers to natural selection, his readers mock him for his fascination for delicate morphological contrivances and co-adaptations – a sign that he was inescapably lured to finality. Some even suggested that his system was a “theodicy”. In the history of Darwinism, the Orchids reveal “another” quite unexpected and heterodox Darwin: freed from the hypothetical fancies of the Origin, and even suggesting a new kind of physico-theology.     

Darwin and Dostoyevsky: twins

The Russian poet Fyodor Dostoyevsky published an insightful treatise on human nature in his novel ‘The Brothers Karamazov'' in 1880. His account of humanity may offer as much insight into human nature for scientists as Darwin''s The Descent of Man.Late in the nineteenth century, Charles Darwin (1809–1882) and Fyodor Dostoyevsky (1821–1881) published accounts of their investigation of humankind. Darwin did so in 1871 in his book The Descent of Man, Dostoyevsky in 1880 in the parable of The Grand Inquisitor in his book The Brothers Karamazov. Last year we celebrated Darwin''s anniversary; for biologists, 2010—the 130th anniversary of Dostoyevsky''s book—might have been the year of Dostoyevsky.Dostoyevsky was familiar with Darwin''s doctrine and he was willing to admit “man''s descent from the ape”. An orthodox Christian, he put this sentiment in religious terms: “It does not really matter what man''s origins are, the Bible does not explain how God moulded him out of clay or carved him out of stone. Yet, he saw a difference between humans and animals: humans have a soul.The philosopher Nikolay Berdyayev noticed: “[Dostoyevsky] concealed nothing, and that''s why he could make astonishing discoveries. In the fate of his heroes he relates his own destiny, in their doubts he reveals his vacillations, in their ambiguity his self-splitting, in their criminal experience the secret crimes of his spirit.”The Grand Inquisitor can be read as Dostoyevsky''s treatise on human nature. In the tale, Jesus Christ revisits Earth during the period of the Inquisition and is arrested by the Church and sentenced to death. The Grand Inquisitor comes to visit Jesus in his prison cell to argue with him about their conceptions of human nature. He explains that humankind needs to be ruled to be happy and that the true freedom Jesus offered doomed humanity to suffering and unhappiness. Dostoyevsky''s superposition of these two points of view on humankind reminds us of the principle of complementarity, by which the physicist Niels Bohr attempted to account for the particle-wave duality of quantum physics.Dostoyevsky conceives of humans as complex, contradictory and inconsistent creatures. Humans perceive personal liberty as a burden and are willing to barter for it, as the Grand Inquisitor explained to Christ, for “miracle, mystery, and authority”. In addition, “the mystery of human being does not only rest in the desire to live, but in the problem: for what should one live at all?” We might say that these faculties make Homo sapiens a religious species. Not in the sense of believing in gods or a god, but in the sense of the Latin word religare, which means to bind, connect or enfold. Humans are mythophilic animals, driven by a need to find a complete explanation for events in terms of intentions and purposes.Research into the neurological bases of imagination, transcendence, metaphorability, art and religion, as well as moral behaviour and judgement (Trimble, 2007) is consistent with Dostoyevsky''s views. It has identified areas of the brain that have been labelled as the ‘god module'' or ‘god spot'' (Alper, 2001). These areas represent a new stratum of evolutionary complexity, an emergence specific to the human species. Their mental translations might be tentatively designated as the Darwinian soul, anchored in the material substrate and neither immortal nor cosmic. As consciousness and volition have become legitimate subjects of neuroscience (Baars, 2003), the Darwinian soul, and with it spirituality, seems to be ripe for scientific inquiry: the quest for meaning, creation and perception of metaphors, the experience of the trinity of Truth, Good and Beauty, the capacity for complex feelings that Immanuel Kant called sublimity, the thrill of humour and play, the power of empathy and the follies of boundless love or hate. Secularization does not erase the superstructure of spirituality: it is reflected, however queer it might seem, in the hypertrophy of the entertainment industry and also, more gloomily, in spiritual conflicts on a global scale.Dostoyevsky''s views on the human soul might be closer to those of Alfred Russel Wallace, who believed that an unknown force directed evolution towards an advanced organization. We can identify this ‘force'' as the second law of thermodynamics (Sharma & Annila, 2007). By moving evolving systems ever farther away from equilibrium, the second law eventually became the Creator of the ‘Neuronal God''.Christ, in the parable of the Grand Inquisitor, might be conceived of as a symbol of the truth outside the human world. Christ was listening to the assertions and questions of his interlocutor, but did not say a single word. His silence is essential to the parable.Similarly, the cosmos, to which humanity has been addressing its questions and predications, remains silent. By science, we increase knowledge only by tiny increments. The ‘god modules'' of our brains, unsatisfied and impatient, have hastily provided the full truth, deposited in the Holy Scripture. There are at least three books claiming to contain the revealed and hence unquestionable truth: the Judaic Torah, Christian Bible and Muslim Qur''an. A dogma of genocentrism in biology might offer an additional Scripture: the sequence of DNA in the genomes.Dostoyevsky''s legacy may suggest an amendment to the UN Charter. We, united humankind, solemnly declare: No truth has ever been revealed to us; we respect and tolerate each other in our independent searching and erring.     

The Darwin of pangenesis

The Darwin of pangenesis is very much another Darwin. Pangenesis is Darwin's comprehensive theory of generation, his theory about all sexual and asexual modes of reproduction and growth. He never explicitly integrated pangenesis with his theory of natural selection. He first formulated pangenesis in the 1840s and integrated it with the physiology, including the cytology, of that era. It was, therefore, not consilient with the newer cytology of the 1860s when he published it in 1868. By reflecting on the role of pangenesis in Darwin's life and work, we can learn to take a wider view of his most general theorising about animal and plant life.     

Cross- and self-fertilization of plants

This essay considers Charles Darwin's late work, Cross- and Self-Fertilization of Plants, locating it in the overall context of Darwin's thought and ideas. It is shown how it is part of a long-term interest in the purpose of sexuality, and how it complements Darwin's earlier book on the fertilization of orchids. It is concluded, however, that Darwin had no full solution to his problem.     

On Darwin's palaeontology in The Origin of Species

I investigate the role of palaeontology within Darwin's works through an analysis of the two chapters of The Origin of Species most especially devoted to this science. Palaeontology may occupy several places within the structure of the argumentative logic of Darwinism, but these places have remained to some extent ancillary. Indeed, palaeontology could well document evolutionary patterns, showing the actual occurrence of evolution as a general “historical fact”, but it was poorly adapted to demonstrate the main point of Darwinism: the actual evolutionary process: natural selection acting among individuals. I also show, in agreement with Gould, that Darwin had great confidence in the ultimate ability of palaeontology to support his theory, and that in interpreting palaeontological evidence, he expressed a vision of natural selection much wider and more eclectic than that which has generally been ascribed to him.     

PERSPECTIVE: DARWIN,PROGRESS, AND ECONOMIC PRINCIPLES

Darwin's concept of progress was an economic one, with the accumulation of adaptations having a strong analogy to technological innovations. This interpretation makes it easier to understand his differences with Lamarck and others whose views he considered metaphysically objectionable. It also clarifies his relationship to Malthus and such features of his theory as the episodic nature of evolutionary change. His position is consistent with modern theories of biotic invasions and long-term evolutionary trends. It also accords well with current efforts to produce a synthesis between biology and economics.     

Darwin,the amphibians,and the natural selection

A critical review of Darwin's publications shows that he did not dissert much about amphibians, in comparison with the other tetrapods. However, in “A Naturalist's Voyage round the World”, Darwin described for the first time several amphibian species and was surprised by their peculiar way of life, terrestrial or euryhaline. These amphibian observations around the world led Darwin to discuss evolutionnary notions, like developmental heterochronies or evolving convergences, and later to illustrate his famous natural selection theory. This is confirmed, for example, by the publication of “On the Origin of Species” where Darwin ironically questioned creation theory, trying to explain the absence of amphibians on oceanic islands. Lamarck also considered amphibians as relevant material to illustrate his theory of acquired character heredity. These historical uses of lissamphibians as evolutionary models have been mostly realized before any amphibian fossil discovery, i.e. out of a palaeontological context.     

Haeckel und das Schöne in der Natur

The influential evolutionary biologist Ernst Haeckel (1834–1919), well‐known as a highly disputatious defender of Darwin's work, sought to unite science, philosophy, ethics and art in an all‐embracing world view that he called ?monism“. In this essay his ideas and reflections on aesthetics in nature and their application are reviewed. According to Haeckel, art should be based on motifs that are to be found in the diversity of life forms, which represent, in his opinion, the highest imagineable specification in aesthetics. Beauty in nature should open men's way to nature, and man must not place himself in opposition to nature. Haeckel himself, who was also a gifted artist, helped find the way to such an attitude by publishing thousands of drawings of organisms, mostly microscopically small marine species. His illustrations made organismic structures accessible that a broader public was previously almost unaware of. With these representations he was most influential in almost all areas of art around the turn of the century, including architecture, interior design, painting, glass art and furniture design.     

Darwin the geologist: Between Lyell and von Buch

Upon returning from his voyage on the Beagle, Darwin prepared reports of his geological observations. Together, these reveal Darwin's approach to reasoning about geology. Darwin argued that successive terraces prove a very gradual elevation of the coast that lagoon islands show a reciprocal sinking of the oceanic floor. Hence, Darwin reinforced Lyell's uniformitarian, or steady state theory. Unlike lagoon islands, the movement of erratic boulders onto the plains is evidence of forces, which do not now exist. Darwin and Lyell attributed this movement to floating icebergs. However, mountain formation remained difficult for them to explain with reference to contemporary causes. Lyell discovered uplifts in Scandinavia, which resulted from epirogenesis, whereas mountain formation is an orogenesis, which involves both folding and uplift. Darwin was more impressed by uplift than by folds. However, when in Cordillera he saw strata overturned by masses of injected rock, proving successive periods of violence, Darwin took a position, which was closer to the plutonic theories of von Buch and Humboldt than it was to Lyell's uniformitarian views.     

Darwin's apes and “savages”

Since his visit to Tierra del Fuego in the 1830s, Darwin had been fascinated by the “savages” that succeeded in surviving on such a “broken beach”, and because they were certainly similar in behaviour to our ancestors. However, he was also fascinated by baboons’ behaviour, according to Brehm's accounts: hamadryas baboons showed a strong altruism to the point of risking their own lives in order to save their infants from attack by dogs. In 1871, he mentions he would rather have descended from brave baboons than from “savages”, considered egoistic. We study the two sources of these ideas and try to show how Darwin's comparative reflections on apes and “savages” made him the first evolutionist anthropologist.