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For a long time neurons have blinded neuroscientists with their dazzling array of impulses and synapses. In thrall to these complex neural networks neuroscientists have often concentrated almost exclusively on neurons while the role of the macroglia – the oligodendrocytes and astrocytes – has remained in the shadows. We hope that this book will present a more integrated view of the relationship between oligodendrocytes and neurons and the critical role both cell types play in the central nervous system (CNS).
We aim to set out major aspects of the biology of the oligodendrocyte – a very large, very complex and dynamic cell – highlighting its extraordinarily unique organization and its multiple functions. For example, each oligodendrocyte can produce a plethora of up to 50 elongated paddle-like processes, each of which spirals around an internode of a different CNS axon. This spiraling process forms the compacted myelin lamellae and the associated uncompacted inner mesaxon, lateral paranodal regions and the outer mesaxons so often overlooked. The metabolic requirements and maintenance of such an elaborate organization of membranes depend on the uncompacted and compacted myelin compartments remaining in continuity. This continuity is achieved via the transverse processes and Schmidt–Lanterman incisures and ensures that vital cytoplasmic components have access to the compact myelin membranes. The orchestration of oligodendrocyte interaction with the neurons and other CNS cells is dependent on the precision of developmental processes including cell division, differentiation and migration to exact locations.
If one desired to throw new light on the effect of disease, or injury, and of the process of healing in the brain, the best hope lay in the study of the non-nervous cells.
No Man Alone, Wilder Penfield, 1977 (Gill and Binder, 2007)
For the past 160 or so years the cells of the nervous system have been divided into two main categories: neurons and glia (Kettenmann and Verkhratsky, 2008). Prior to this, ever since the first image of a neuron was published in 1836 by Gabriel Valentin, the nerve cell had been in a class of its own (Lopez-Munoz et al., 2006). Some 20 years later in 1856 the term neuroglia was introduced by the German physician Rudolph Virchow. Virchow, also known as the “Pope of pathology” (Kettenmann and Ransom, 2005; Magner, 2002), described a “connective substance … in which nervous system elements are embedded” and referred to it as “nervenkitt” (or nerve putty). This description led to the use of the term “neuroglia,” which derives from archaic Greek, meaning something sticky or clammy. The notion that neuroglia were there merely as neural putty was treated with the reverence usually reserved for a bona fide papal encyclical and as such neuroglia remained sidelined for decades to come. Even though Virchow was responsible for the term neuroglia coming into use, at this stage he did not recognize that it was made up of cells rather than an acellular connective tissue.
Traditionally, oligodendrocytes have been assumed to play a minor supporting role in the central nervous system and their importance has generally been overlooked. For the first time, this book provides a dedicated review of all of the major aspects of oligodendrocyte biology, including development, organization, genetics, and immunobiology. Later chapters emphasize the importance of this underestimated cell to the mammalian central nervous system by exploring the role of myelin synthesis and maintenance in neural disease and repair. Particular attention is paid to multiple sclerosis (MS), arguably the prime example of an acquired demyelinating disease, with detailed examinations of the current concepts regarding demyelination, oligodendroglial damage, and remyelination in MS lesions.
Schwann cells are a diverse group of cells formed from neural crest cells. They are essential components of the peripheral nerves of both vertebrate and invertebrate nervous systems. The diversity of Schwann cell subsets and function is seen in those Schwann cells that form myelin - that uniquely specialised part of the plasma membrane that spirals around axonal lengths to myelinate the peripheral nerves. The Biology of Schwann Cells concentrates on the cells of mammals and in particular humans. It covers the distinction between compact and non-compact myelin in depth, along with the perisynaptic cells which form the partnership between nerve terminals and muscle fibre. Developmental aspects are discussed alongside differentiation, and the genetics of the cells in health and disease. With chapters from world-renowned experts, this book is aimed at postgraduates and researchers in neuroscience and neurology and anyone involved in the study of peripheral nerves.
It is now over 200 years since Theodore Schwann first described the cell which bears his name. Such early descriptions of nervous system components were done without the powerful microscopes we have today, yet Schwann and Ramon Y. Cajal made foundation observations which still stand. Cajal's papers, especially, show the power of careful observation, an essential element of good science.
The Schwann cell has been historically underrated and poorly understood. In particular, the myelin-forming Schwann cells or their myelin are still often referred to as a simple ‘sheath’ for the neuron. However, Schwann cells in all their complexity form essential partnerships with neurons, and muscles. This is of particular relevance in the case of the myelin-forming Schwann cell, an enormous cell that expresses unique molecules and complex relationships related to maintenance of the compact and non-compact myelin regions of its plasma membrane. Schwann cells have other complex interactions, not least of which are found where nerve terminals and muscle fibres form the tripartite synapse in association with the perisynaptic Schwann cells. There are also the poorly understood satellite cells that surround the dorsal root ganglion nerve cell bodies, and of course the complexity of non-myelinated Schwann cells and their axonal associations.
It may be that the histopathological prominence of abnormalities of compact myelin has focussed research on this region of the Schwann cell.
The Schwann cell is named in honour of the German physiologist Theodor Schwann (1810–1882, Figure 1.1) who is now acknowledged as the founder of modern histology. In addition to describing the Schwann cell, he made numerous contributions to the fields of biology, physiology and histology – not least as one of the instigators and main advocates of cell theory. The cell theory defined the cell as the base unit of all living organisms, and had great influence on the study of both plants and animals. The cell theory was radical for the time and irrevocably discredited Vitalism, the mainstream belief that life was attributed to a vital force. Among other things, Schwann is known for recognising that the crystals seen during fermentation, first reported by Leeuwenhoek in 1680, were in fact living organisms; although it was not until Pasteur in 1878 wrote to Schwann acknowledging this observation that Schwann's finding was accepted. In fact, Pasteur's germ theory stems from Schwann's work in which he showed that microorganisms were required for the putrefaction of meat.
Schwann spent his undergraduate years at the University of Bonn and then the equivalent of postgraduate study in Wuerzburg and Berlin. Schwann was appointed Professor of Anatomy at Louvain in 1839. In 1848 he moved to the Chair of Anatomy in Liege. In a biography of Schwann (Causey 1960), Causey reported that he avoided the strife of scientific controversy and appears to have risen above petty jealousies.
Patricia Armati, Associate Professor in the School of Biological Sciences and the Nerve Research Foundation, University of Sydney, Australia,
Chris Dickman, Professor in Ecology in the Institute of Wildlife Research School of Biological Sciences, University of Sydney, Australia,
Ian Hume, Professor in the School of Biological Sciences University of Sydney, Australia
It is now more than 30 years since Hugh Tyndale-Biscoe's student text Life of Marsupials (1973) was published, and almost 30 years since the first compendia on marsupial biology appeared. Both bore the same title, The Biology of Marsupials, and were edited by B. Stonehouse and D. Gilmour (1977) and D. Hunsaker II (1977). Since then numerous more specialised books on various aspects of marsupial biology have appeared. However, with the exception of a new edition of Life of Marsupials (Tyndale-Biscoe 2005) that appeared while the current book was in production, none has the breadth of the earlier books. The closest is Marsupial Biology: Recent Research, New Perspectives, edited by N. R. Saunders and L. A. Hinds (1997). There is thus a need for a resource book that covers the many facets of marsupial biology, many of which are unique to the mammalian subclass Marsupialia.
In Marsupials we have harnessed the collective knowledge and wisdom of a select group of colleagues from the Americas and Australia. The result is a collection of essays that cover marsupials from their beginnings and subsequent evolution, through their genetics, anatomy, physiology, ecology and behaviour, to conservation management concerns. Each chapter stands as a view into the marsupial world by its author(s). Although all chapters have been independently reviewed, then edited for consistency and cross-referencing to other chapters, the original style has been retained in all cases.