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areas of the brain instead of a result of the presentation of a stimulus. This interpretation represents a major advance beyond purely sensory theories of consciousness. It correctly grants attention (the brains ability to select particular events for special processing) a major role in structuring our conscious experience. Koch also stresses the adaptive value of such conscious experience in human evolution. Consciousness allows us to plan our actions better, by extracting the crucial information we need about the external world. Some simple actions, such as reaching out to intercept an object coming toward us, occur too fast for executive decisions. Accordingly, we can respond to such stimuli without consciousness. Koch calls these specialized sensory-motor processes zombie agents, which are not the same as the zombies philosophers use in their discussions of the nature of consciousness. The mental ability of humans goes far beyond the limited powers of these zombie agents, and its higher levels may require consciousness. What Koch cannot tell us is how the control of action constrains and controls conscious experience: this will emerge as a key question in neuroscience over the next decade. Qualia are perhaps the greatest problem for the NCC approach. Why does it feel like something when a coalition of neurons fires? Why should neural activity produce any kind of feeling at all? Koch acknowledges that he sidesteps these questions. An NCC model that combines visual analysis and executive control faces the problem of why the content of our awareness is often simple attributes (such as color or pitch) rather than whole objects or executive decisions. But Koch is correct in suggesting that some useful scientific progress can be made without being tortured by the philosophical problems of qualia. For example, although studies of bistable percepts might not account for qualia, they may at least indicate how qualia are generated. The books presentation has several strong points. Koch begins with an excellent introductory chapter on the problem of consciousness in neuroscience, which includes short summaries of the various answers that have been proposed. He ends each chapter with a very helpful recapitulation, allowing readers to check their intellectual progress. Although not lavish, the illustrations are pertinent and clear. The book is least successful in its middle chapters, which seem overly long and in places unsatisfactory. There, Koch offers readable summaries of important recent work in visual cognitive

Seeing Through the Steam of Consciousness

Patrick Haggard

The reviewer is at the Institute of Cognitive Neuroscience and Department of Psychology, University College London, London WC1N 3AR, UK. E-mail: p.haggard@ucl.ac.uk

Exotic Landscape (Henri Rousseau, 1910).

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CREDIT: SUZUKI COLLECTION, TOKYO, JAPAN/PETER WILLI/BRIDGEMAN ART LIBRARY

onsciousness has often been described as one of the great scientific challenges of modern times. Our existence is characterized by the experience of what we see, the feel of what we touch, and the internal sensations that are linked to emotions. The philosophical tradition of dualism treats The Quest for these conscious mental Consciousness events as quite separate A Neurobiological from the material world, Approach including the physical Christof Koch matter of the brain. Most Roberts, Englewood, modern neuroscientists CO, 2004. 447 pp. $45, 29.99. ISBN 0- reject dualism and argue that conscious states 9747077-0-8. must arise from physical events in the brain, notably the electrical impulses in neurons of the cerebral cortex. The Quest for Consciousness does not really explain why patterns of electrical activity can produce consciousness. Instead, by summarizing over two decades of research on the brain mechanisms of visual perception, Christof Koch aims to give a detailed account of which patterns of brain activity produce conscious experience. His exciting and compelling book provides a detailed, serious attempt at a scientific explanation of consciousness. At the heart of the authors approach is the search for neuronal correlates of consciousness (NCCs). Koch, a cognitive scientist at Caltech, defines these as the minimal neuronal events jointly sufficient for a specific conscious percept. That is, if a specific coalition of neurons is electrically active, then this produces the specific conscious experience of seeing Bill Clinton, having a splitting headache, or whatever. Some strong evidence for this view has been obtained by monitoring neuronal responses during the alternating bistable percepts that may occur when different images are presented to each of a subjects eyes. Thus if a monkey is trained to respond to indicate which of two stimuli it currently sees, some visual neurons will fire specifically during one percept but not the other. The NCC idea is rooted in the classical findings of tuned or specialized neurons in the visual cortex,

neurons that are activated by particular shapes, colors, or directions of motions. Koch proposes some additional neural processes, over and above feature detection, which contribute to conscious experience. Broadly speaking, these involve interactions between primary visual cortex and several nonprimary cortical areas. Kochs discussion of these interactions forms the most innovative and interesting part of the book. For example, Koch argues that activity in primary visual cortex is not sufficient for consciousness. (That may be an important difference from other sensory systems, because the elegant research of Ranulfo Romo suggests that artificial stimulation of primary somatosensory cortex may generate a genuine tactile experience of vibration.) Instead, Koch links conscious experience to a traditional hierarchical view of visual object analysis in the cortex. A chain of secondary cortical areas shows increasing specificity for particular classes of stimuli, ending with explicit representations of complex stimuli such as faces by neurons in the inferotemporal cortex. These neurons are effectively NCCs. On Kochs view, we should be conscious of objects and interpretations, rather than having raw, primary experiences (what philosophers have called qualia). Another key claim of Kochs position is that consciousness is something we control, rather than something the visual world forces upon us. Koch emphasizes that feedback from cortical areas associated with planning and action into perceptual areas plays a crucial role in generating conscious experience. On this view, consciousness becomes an executive decision by the frontal

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BOOKS ET AL.
searcher at the Marine Biological Laboratory, Woods Hole) have tackled the task of compiling what we currently know about the islands environments, ecology, and biota. Working with some 300 contributors, 70 of whom are Malagasy researchers, they have produced a hefty volume comprising 14 chapters, with over 250 individually written sections and thousands of references, and running nearly 1800 pages. Popular knowledge associates Madagascar with living lemurs, tenrecs, and hissing cockroaches as well as recently extinct giant lemurs and 400-kg elephant birds. The Natural History of Madagascar explodes that narrow view with a wealth of detail. Lists appear everywhere throughout the volumeto name a few: a taxonomic list of the modern bird fauna, the collection BIODIVERSITY localities and habitat for each of the known 685 species of Malagasy gastropods, presence data on each of the almost 500 papilionid legumes found in or around parks and protected areas, and a checklist of the ferns and their allies. Despite a long history of scientific exploration in Madagascar (which began in the mid-17th century), much of the John G. Robinson information in the book was collected only adagascar is simply more different in the last few decades. Inevitably, there are than most places. Over 80% of the many omissions, but the coverage is relaislands 12,000 described plant tively complete and the descriptions are ofspecies and 70% of its butterfly ten presented down to the species are endemic. Of the species level. The Natural History 141 species of freshwater fish Although the middle seven of Madagascar known from the island, 91 are chapters, the bulk of the book, Steven M. Goodman and found nowhere else. Only 2 of are organized taxonomically, Jonathan P. Benstead, Eds. the islands 203 described this is not a dry litany of biospecies of tiger beetles are University of Chicago Press, logical diversity. Six introducnonendemic. All of the 101 na- Chicago, 2003. 1731 pp. tory chapters place Madagas$85, 59.50. ISBN 0-226tive species of terrestrial mam- 30306-3. car in its geologic, climatic, mals are restricted to the island ecological, and anthropologior the nearby Comoro Islands. cal contexts, and the volume The evolution responsible for ends with a consideration of this rich endemic biota has been proceeding the conservation challenge. Throughout, spein insular isolation for a long time. What was cialists were clearly encouraged to provide to become Madagascar separated from the their own perspective. Thus interwoven with rest of Pangea some 150 million years ago more taxonomic sections are others on such and then split off from what would become topics as the pollination ecology of plant the Indian subcontinent some 60 million communities in dry forests, the occurrence of years later. alkaloids and phenolics in forests as well as Steven Goodman (a Field Museum biol- the responses by lemurs to these plant comogist whose name is almost synonymous pounds, and the consequences of forest fragwith natural history in Madagascar) and mentation for the herpetofauna. The reader Jonathan Benstead (a postdoctoral re- can ponder natural history puzzles: Why Takhtajania perrieri, a small tree in the Winteraceae (which today only occurs addiThe reviewer is at the Wildlife Conservation Society, tionally in South America and Pacific is2300 Southern Boulevard, Bronx, NY 104601099, lands), apparently has no seed disperser and USA. E-mail: jrobinson@wcs.org neuroscience, and he also tackles a large number of other topics ranging from shortterm memory to split-brain patients. All these topics are loosely related to consciousness. But only in very different ways, and they are not drawn together with clear structuring principles. The authors approach to consciousness may be overly ambitious at these points, and he sometimes abandons the rigor that the combination of visual psychophysics and recordings from single neurons in monkeys has offered him. His central account might emerge more strongly if the book were pruned or read selectively. Nonetheless, Kochs writing style clear, colloquial, and helpfulis very successful. As a result, the book usefully fills a gap in the coverage of consciousness. It is more scientific and intellectually rigorous than most popular accounts as well as much more accessible than the primary research literature. The Quest for Consciousness is a brave attempt to fuse the best scientific thinking with one of the central aspects of human existence. seedlings typically cluster around the parent plant. Why the shrew tenrec Microgale longicaudata has a very long prehensile tail and elongate hindfeet and digits. Anyone who likes to rummage in the attic of natural history facts will find the book a treat. Did you know, for instance, that in the 19th century, Nephila spiders were raised in special gardens for silk production? This compilation of data will be invaluable to both specialists in particular taxa and those with general interests in Madagascar. But more than anything, the volume highlights the importance of species as a focus for conservation. Conservationists are justifiably concerned with biodiversity writ largegenes, species, communities, biotic processes, ecosystems, and biomes. Ecosystem management is our current mantra. We are preoccupied with the maintenance of ecosystem function and the provision of ecosystem services for resources, basic lifesupport systems, and human enjoyment of nature. Nevertheless, species are the building blocks for biodiversity, and species both provide the tangible resources and underlie the higher level processes. By monitoring the effect of conservation programs on species, one can evaluate management programs and land use changes. Two decades ago, a volume edited by Daniel Janzen [Costa Rican Natural History (University of Chicago Press, 1983)] celebrated the species diversity of another tropical country. It has since supported the flowering of the biological and conservation traditions there. One hopes that this book will do the same for Madagascar. Yet all too easily could this diversity be lost. There is a thread of panic woven through the volume. In commenting on freshwater fishes, John Sparks and Melanie Stiassny state simply that neither of us has witnessed anything surpassing the degree and extent of environmental degradation in Madagascar. It may seem as if so much of the cataloging is just to have a record of what might be lost. But the books last chapter offers a more optimistic conclusion. Madagascar is universally recognized as a huge conservation priority, and the island is receiving ever-increasing investments, both individual and institutional, in conservation. The countrys new president, Marc Ravalomanana, has made a personal commitment to conserving Madagascars unique (a word not used lightly) biological diversity. By dint of all its numbers, The Natural History of Madagascar makes the case for what we need to save.

CREDIT: DETAIL FROM THE WWF-MADAGASCARS POSTER FAUNAL BIODIVERSITY OF MADAGASCAR, BY HARALD SCHTZ/COURTESY UNIVERSITY OF CHICAGO PRESS

An Island of Evolutionary Exuberance

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