Peter Holland,homeobox genes,and the developmental basis of animal diversity

In 1867 Alexander Kowalevsky published an account of the development of the cephalochordate Amphioxus lanceolatus (now known as Branchiostoma lanceolatum) (Kowalevsky, 1867). Together with his study of the development of urochordates (Kowalevsky, 1866; 1871), this introduced a new way of thinking about the relationship between the evolution and development of animals, and established the basis for long-standing theories of the evolutionary origin of vertebrates. Some one hundred and fifty years later, cephalochordates and urochordates are again in the spotlight, as molecular biology and genome sequencing promise further revelations about the origin of vertebrates. The work of the 2006 Kowalevsky Medal winner, Peter Holland (Fig. 1), has played a central role in their reinstatement (see Mikhailov and Gilbert (2002) for more details of the history of the Kowalevsky Medal). Here, I profile Peter Holland’s contribution to the rebirth of Evolutionary Developmental Biology in general and the study of homeobox genes and vertebrate origins in particular.     

From development to evolution: the re-establishment of the "Alexander Kowalevsky Medal"

The Saint Petersburg Society of Naturalists has reinstated the Alexander O. Kowalevsky Medal. This article announces the winners of the first medals and briefly reviews the achievements of A.O. Kowalevsky, the Russian comparative embryologist whose studies on amphioxus, tunicates and germ layer homologies pioneered evolutionary embryology and confirmed the evolutionary continuity between invertebrates and vertebrates. In re-establishing this international award, the Society is pleased to recognize both the present awardees and the memory of Kowalevsky, whose work pointed to that we now call evolutionary developmental biology.     

Peter Holland, homeobox genes and the developmental basis of animal diversity

In 1867 Alexander Kowalevsky published an account of the development of the cephalochordate Amphioxus lanceolatus (now known as Branchiostoma lanceolatum) (Kowalevsky, 1867). Together with his study of the development of urochordates (Kowalevsky, 1866; 1871), this introduced a new way of thinking about the relationship between the evolution and development of animals and established the basis for long-standing theories of the evolutionary origin of vertebrates. Some one hundred and fifty years later, cephalochordates and urochordates are again in the spotlight, as molecular biology and genome sequencing promise further revelations about the origin of vertebrates. The work of the 2006 Kowalevsky Medal winner, Peter Holland has played a central role in their reinstatement. Here, I profile Peter Holland's contribution to the rebirth of Evolutionary Developmental Biology in general and the study of homeobox genes and vertebrate origins in particular.     

Peter Holland, homeobox genes, and the developmental basis of animal diversity

In 1867 Alexander Kowalevsky published an account of the development of the cephalochordate Amphioxus lanceolatus (now known as Branchiostoma lanceolatum) (Kowalevsky, 1867). Together with his study of the development of urochordates (Kowalevsky, 1866; 1871), this introduced a new way of thinking about the relationship between the evolution and development of animals, and established the basis for longstanding theories of the evolutionary origin of vertebrates. Some one hundred and fifty years later, cephalochordates and urochordates are again in the spotlight, as molecular biology and genome sequencing promise further revelations about the origin of vertebrates. The work of the 2006 Kowalevsky Medal winner, Peter Holland (Fig. 1), has played a central role in their reinstatement (see Mikhailov and Gilbert (2002) for more details of the history of the Kowalevsky Medal). Here, I profile Peter Holland's contribution to the rebirth of Evolutionary Developmental Biology in general and the study of homeobox genes and vertebrate origins in particular.     

Kowalevsky, comparative evolutionary embryology, and the intellectual lineage of evo-devo

Alexander Kowalevsky was one of the most significant 19th century biologists working at the intersection of evolution and embryology. The reinstatement of the Alexander Kowalevsky Medal by the St. Petersburg Society of Naturalists for outstanding contributions to understanding evolutionary relationships in the animal kingdom, evolutionary developmental biology, and comparative zoology is timely now that Evo-devo has emerged as a major research discipline in contemporary biology. Consideration of the intellectual lineage of comparative evolutionary embryology explicitly forces a reconsideration of some current conceptions of the modern emergence of Evo-devo, which has tended to exist in the shadow of experimental embryology throughout the 20th century, especially with respect to the recent success of developmental biology and developmental genetics. In particular we advocate a sharper distinction between the heritage of problems and the heritage of tools for contemporary Evo-devo. We provide brief overviews of the work of N. J. Berrill and D. T. Anderson to illustrate comparative evolutionary embryology in the 20th century, which provides an appropriate contextualization for a conceptual review of our research on the sea urchin genus Heliocidaris over the past two decades. We conclude that keeping research questions rather than experimental capabilities at the forefront of Evo-devo may be an antidote to any repeat of the stagnation experienced by the first group of evolutionary developmental biologists over one hundred years ago and acknowledges Kowalevsky's legacy in evolutionary embryology.     

On some historical and theoretical foundations of the concept of chordates

The concept of chordates arose from the alliance between embryology and evolution in the second half of the nineteenth century, as a result of a theoretical elaboration on Kowalevsky’s discoveries about some fundamental similarities between the ontogeny of the lancelet, a putative primitive fish, and that of ascidians, then classified as molluscs. Carrying out his embryological studies in the light of Darwin’s theory and von Baer’s account of the germ layers, Kowalevsky was influenced by the German tradition of idealistic morphology that was concerned with transformations driven by laws of form, rather than with a gradual evolution occurring by means of variation, selection and adaptation. In agreement with this tradition, Kowalevsky interpreted the vertebrate-like structures of the ascidian larva according to von Kölliker’s model of heterogeneous generation. Then, he asserted the homology of the germ layers and their derivatives in different types of animals and suggested a common descent of annelids and vertebrates, in agreement with Saint-Hilaire’s hypothesis of the unity of composition of body plans, but in contrast with Haeckel’s idea of the Chordonia (chordates). In The Descent of Man Darwin quoted Kowalevsky’s discoveries, but accepted Haeckel’s interpretation of the ascidian embryology within the frame of a monophyletic tree of life that was produced by the fundamental biogenetic law. Joining embryology to evolution in the light of idealistic morphology, the biogenetic law turned out to be instrumental in bringing forth different evolutionary hypotheses: it was used by Haeckel and Darwin to link vertebrates to invertebrates by means of the concept of chordates, and by Kowalevsky to corroborate the annelid theory of the origin of vertebrates. Yet, there was still another interpretation of Kowalevsky’s discoveries. As an adherent to empiricism and to Cuvier’s theory of types, von Baer asserted that these discoveries did not prove convincingly a dorsal position of the nervous system in the ascidian tadpole larva; hence, they could not support a homology between different animal types suggesting a kinship between ascidians and vertebrates.     

Michael Akam and the rise of evolutionary developmental biology

Michael Akam has been awarded the 2007 Kowalevsky medal for his many research accomplishments in the area of evolutionary developmental biology. We highlight three tributaries of Michael’s contribution to evolutionary developmental biology. First, he has made major contributions to our understanding of development of the fruit fly, Drosophila melanogaster. Second, he has maintained a consistent focus on several key problems in evolutionary developmental biology, including the evolving role of Hox genes in arthropods and, more recently, the evolution of segmentation mechanisms. Third, Michael has written a series of influential reviews that have integrated progress in developmental biology into an evolutionary perspective. Michael has also made a large impact on the field through his effective mentorship style, his selfless promotion of younger colleagues, and his leadership of the University Museum of Zoology at Cambridge and the European community of evolutionary developmental biologist.     

Russian comparative embryology takes form: a conceptual metamorphosis toward “evo‐devo”

This essay recapitulates major paths followed by the Russian tradition of what we refer to today as evolutionary developmental biology (“evo‐devo”). The article addresses several questions regarding the conceptual history of evolutionary embryological thought in its particularly Russian perspective: (1) the assertion by the St. Petersburg academician Wolff regarding the possible connections between environmental modifications during morphogenesis and the “transformation” of species, (2) the discovery of shared “principles” underlying animal development by von Baer, (3) the experimental expression of Baer's principles by Kowalevsky and Mechnikoff, (4) Severtsov's theory of phylembryogenesis, (5) Filatov's approach to the study of evolution using comparative “developmental mechanics”, and (6) Shmalgausen's concept of “stabilizing” selection as an attempt to elucidate the evolution of developmental mechanisms. The focus on comparative evolutionary embryology, which was established by Kowalevsky and Mechnikoff, still continues to be popular in present‐day “evo‐devo” research in Russia.     

The meteoric rise of regulated intracellular proteolysis

It is often the case in biology that research into breaking things down lags behind research into synthesizing them, and this is certainly true for intracellular proteolysis. Now that we recognize that intracellular proteolysis, triggered by attaching multiple copies of a small protein called ubiquitin to target proteins, is fundamental to life, it is hard to believe that 20 years ago this field was little more than a backwater of biochemistry studied by a handful of laboratories. Among the few were Avram Hershko, Aaron Ciechanover and Alexander Varshavsky, who were recently awarded the Albert Lasker award for basic medical research for discovering the importance of protein degradation in cellular physiology. This Timeline traces how they and their collaborators triggered the rapid movement of ubiquitin-mediated proteolysis to centre stage.     

No evidence for linkage disequilibrium between Bf and GLO in African Negroids

Summary A sample of South African Negroids (n=791) was scored for each individual's Bf and GLO phenotype. (The genes for the Bf and GLO polymorphisms are included in a known cluster of linked genes on chromosome 6.) Following a ²-test the respective two series of alleles were found to be distributed at random, i.e., there was no evidence for a linkage disequilibrium. This result is discussed in terms of the linkage relationships and map distances of the genetic markers involved.Supported by the Deutsche Forschungsgemeinschaft (DFG), Bonn-Bad GodesbergSupported by a grant from the Georg and Agnes Blumenthal-Stiftung, BerlinSupported by a research fellowship (1975/76) awarded by the Alexander von Humboldt-Stiftung, Bonn-Bad Godesberg     

Launching Channels: A New Milestone in the Ion Channel Field

Ion channels underlie a plethora of physiological functions not only in the animal kingdom, but also in plants and microorganisms such as bacteria. Even though we have only known of the existence of channels for about four decades, a PubMed search for channels yields over 120,000 papers, with 40,000 of those appearing in the past five years alone. Even before ion channels had been formally discovered, their existence was hypothesized by Hodgkin and Huxley who were awarded the Nobel Prize in Physiology/Medicine for their work on electrical activity in axons. Subsequent Nobel awards for ion channel electrophysiology techniques (Bert Sakmann and Erwin Neher for Medicine in 1991) and ion channel structure and physiology (Rod MacKinnon and Peter Agre for Chemistry in 2003) underscored the contemporary importance of ion channel research. It is noteworthy that single channel recording is one of the most sensitive techniques in biology – allowing researchers to study the function of a single molecule in its native environment in real time.     

WLADIMIR KOWALEVSKY'S SOURCES OF IDEAS AND THEIR IMPORTANCE FOR HIS WORK AND FOR RUSSIAN EVOLUTIONARY PALAEONTOLOGY

Studies of documents and data about W.O. Kowalevsky's life and work have shed new light on his scientific background. After graduating from the School of Jurisprudence in St. Petersburg, he was first engaged in book publishing, but in 1868 began studies at the Anatomical Institute of the Medical-Surgical Academy, founded by N.I. Pirogov, a famous surgeon and anatomist. Pirogov's ideas were adopted by P.F. Lesshaft (1837–1907), later a prominent anatomist and founder of functional anatomy in Russia. Another teacher was the famous physiologist I.M. Sechenov. Only with the background of this new functional anatomy was Kowalevsky able to link up various data to form his own theories and create classical palaeontological works in the space of two and a half years (1871–1873). His influence on the development of vertebrate palaeontology has been lasting. In Russia and the Soviet Union, Kowalevsky and Lesshaft had such successors as A.P. Pavlov (1854–1929), A.A. Borissiak (1872–1944), N.N. Yakovlev (1870–1966), and many other disciples in younger generations of palaeontologists.     

Reliability of enzyme systems and molecular mechanisms of aging

A stochastic model of animal mortality and ageing is proposed. In a living being there is a "biological clock", constructed from Q kinds of functional elements (genes), The reliability of functioning of these genes determines the biological age of the being. Oxidative processes in the cells are accompanied by random failures of electron transfer enzyme systems, which result in appearance of free superoxide radicals. Failures of electron transfer enzymes and of defence enzymes lead to accumulation of functional damages in the genes of the "clock". In terms of the model experimental data of gerontology (Gompertz's law, Strehler-Mildvan's compensation effect, etc) are easily explained. The value of Q for the man is estimated to be of the order of 10. Some possible mechanisms of action of "geroprotectors" have been discussed.     

Alexander disease: models,mechanisms, and medicine

Alexander disease is a primary disorder of astrocytes caused by gain-of-function mutations in the gene for glial fibrillary acidic protein (GFAP), which lead to protein aggregation and a reactive astrocyte response, with devastating effects on the central nervous system. Over the past two decades since the discovery of GFAP as the culprit, several cellular and animal models have been generated, and much has been learned about underlying mechanisms contributing to the disease. Despite these efforts, many aspects of Alexander disease have remained enigmatic, particularly the initiating events in GFAP accumulation and astrocyte pathology, the relation between astrocyte dysfunction and myelin deficits, and the variability in age of onset and disease severity. More recent work in both old and new models has begun to address these complex questions and identify new therapeutics that finally offer the promise of effective treatment.     

Evolutionary embryology resurrected in Japan with a new molecular basis: Nori Satoh and the history of ascidian studies originating in Kyoto during the 20th century

This article briefly summarizes the scientific contributions of Nori Satoh, the winner of the 2005 edition of the Kowalevsky Medal, to Developmental Biology and especially to Evo-Devo with his 30 years of research on tunicates - a primitive chordate species. His research began with his pure developmental interest in the clock mechanism of cell differentiation and later expanded into various aspects of evolutionary and developmental phenomena. He is not only known as a founder of molecular biology-based tunicate studies, but also for his world-wide service to education and his prestigious publications in international scientific journals.     

Medical housing "lines".

General practitioners are often asked for medical certificates (housing "lines") by applicants for council housing who claim to have medical problems requiring housing priority. The results of a survey by questionnaire showed that general practitioners in Edinburgh do not know how the housing system works and that they seem to overestimate their patients'' chances of obtaining suitable council housing. General practitioners need to know how the housing system works, and communication between general practitioners and housing departments should be improved. A comparison was also made between the number of medical points awarded by a community medicine specialist and a group of general practitioners who had written housing "lines" for their patients. The general practitioners tended to award more points than the specialist. Social priority for housing should be recognised as an independent factor and a new category of top social priority added.     

Warfare and hominid brain evolution.

This paper reviews literature on the evolutionary effects of warfare upon the hominid brain. Alexander &; Tinkle (1968) and Bigelow (1969) are found to be the first to propose that warfare was the principle evolutionary pressure that created the novel substance of the human brain, and that it acted at least from the early Pleistocene. These writers are distinguished from Darwin (1871), Keith (1947) and Wilson (1975) who saw warfare influencing the development of the brain only in historical or near-historical times.The warfare hypothesis of Alexander &; Tinkle is found to be an excellent explanation of the evolution of the human brain, but to be unsatisfactory from a biological viewpoint because they do not explain how warfare evolved in the first place, nor do they attempt to account for the apparent absence of warfare as a behavioral adaptation in species other than some eusocial insects.This author underpins the warfare hypothesis, arguing that it evolved as a necessary consequence of the circumstances of early hominids. Proficient tool use gave domination over predators and opened up new food resources, thereby diminishing two population controls. A population explosion resulted and, at critical densities, when starvation threatened, warfare was the genetically most successful behavioral adaptation. Alternative hypotheses are shown to be inadequate. Finally, the author asks why such an important hypothesis has been ignored for almost a decade.