Behavioral Mechanisms in Evolutionary Ecologyby Leslie A. Real

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Behavioral Mechanisms in Evolutionary Ecology by Leslie A. RealReview by: Rachel NeemsJournal of Animal Ecology, Vol. 64, No. 6 (Nov., 1995), pp. 790-791Published by: British Ecological SocietyStable URL: http://www.jstor.org/stable/5864 .Accessed: 07/05/2014 17:53Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .http://www.jstor.org/page/info/about/policies/terms.jsp .JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range ofcontent in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new formsof scholarship. For more information about JSTOR, please contact support@jstor.org. .British Ecological Society is collaborating with JSTOR to digitize, preserve and extend access to Journal ofAnimal Ecology.http://www.jstor.org This content downloaded from 169.229.32.136 on Wed, 7 May 2014 17:53:07 PMAll use subject to JSTOR Terms and Conditionshttp://www.jstor.org/action/showPublisher?publisherCode=britecohttp://www.jstor.org/stable/5864?origin=JSTOR-pdfhttp://www.jstor.org/page/info/about/policies/terms.jsphttp://www.jstor.org/page/info/about/policies/terms.jspsuch complex systems and with such different scales of time and space that different integrative levels are necessary according to the questions asked. This is not to deny that there are interfaces and overlap between the subdisciplines just as the ecosystem approach often requires understanding of physiology, geochemistry, hydrology, etc. Certainly, some ecol- ogists (and among them probably the most influential ones) have contributed to ecosystem studies as well as to population biology and evolutionary ecology, with the understanding that they were asking different questions at different times. I found that the most constructive suggestion in the book is that expressed in the chapter by L. B. Slobodkin. In his view the problem is not that ecology consists of different inte- grative levels (subdisciplines); rather it is a problem of a too narrow training of ecologists. Given a broad background in natural history, physiology, classi- fication, etc., and in different approaches to ecology, students will gain appreciation of different integrative levels and of ecological complexity and they will be able to approach and solve real problems. Altogether, the book includes several interesting chapters which are worthwhile reading. Strictly speak- ing, most fail to contribute to the stated purpose of the book: they exemplify the interface between population and ecosystem ecology rather than their integration. But they often present interesting approaches to par- ticular ecological problems. The majority of the authors write from an ecosystem perspective; the book would, perhaps, have become more interesting and balanced if a larger number of evolutionary or popu- lation ecologists had also been invited to contribute. TOM FENCHEL Leslie A. Real (Ed.) (1994) Behavioral Mechanisms in Evolutionary Ecology. Pp. 478. University of Chicago Press, Chicago. ?63.95 (cloth), ?23.95 (paperback). ISBN 0-226-70597-8. How often have we heard the plea that geneticists, for example, must talk to ecologists or ethologists converse with physiologists? This new volume of pap- ers edited by Leslie Real preaches the message that all functional biologists, whatever their discipline, should talk to evolutionary ecologists and vice versa. There is a lot of sense in what the authors say. Studying hormones and life histories in tandem, for example, can lead to a far greater understanding than when either is studied in isolation (chapter 14, Ketterson & Van Nolan). The common theme that runs through the book is that all ecological phenomena and patterns of com- such complex systems and with such different scales of time and space that different integrative levels are necessary according to the questions asked. This is not to deny that there are interfaces and overlap between the subdisciplines just as the ecosystem approach often requires understanding of physiology, geochemistry, hydrology, etc. Certainly, some ecol- ogists (and among them probably the most influential ones) have contributed to ecosystem studies as well as to population biology and evolutionary ecology, with the understanding that they were asking different questions at different times. I found that the most constructive suggestion in the book is that expressed in the chapter by L. B. Slobodkin. In his view the problem is not that ecology consists of different inte- grative levels (subdisciplines); rather it is a problem of a too narrow training of ecologists. Given a broad background in natural history, physiology, classi- fication, etc., and in different approaches to ecology, students will gain appreciation of different integrative levels and of ecological complexity and they will be able to approach and solve real problems. Altogether, the book includes several interesting chapters which are worthwhile reading. Strictly speak- ing, most fail to contribute to the stated purpose of the book: they exemplify the interface between population and ecosystem ecology rather than their integration. But they often present interesting approaches to par- ticular ecological problems. The majority of the authors write from an ecosystem perspective; the book would, perhaps, have become more interesting and balanced if a larger number of evolutionary or popu- lation ecologists had also been invited to contribute. TOM FENCHEL Leslie A. Real (Ed.) (1994) Behavioral Mechanisms in Evolutionary Ecology. Pp. 478. University of Chicago Press, Chicago. ?63.95 (cloth), ?23.95 (paperback). ISBN 0-226-70597-8. How often have we heard the plea that geneticists, for example, must talk to ecologists or ethologists converse with physiologists? This new volume of pap- ers edited by Leslie Real preaches the message that all functional biologists, whatever their discipline, should talk to evolutionary ecologists and vice versa. There is a lot of sense in what the authors say. Studying hormones and life histories in tandem, for example, can lead to a far greater understanding than when either is studied in isolation (chapter 14, Ketterson & Van Nolan). The common theme that runs through the book is that all ecological phenomena and patterns of com- munity can be viewed as immediate consequences of an individual's behaviour. Furthermore, to fully understand the behaviour one must look at the munity can be viewed as immediate consequences of an individual's behaviour. Furthermore, to fully understand the behaviour one must look at the internal physiological and psychological processes in conjunction with the ecological and evolutionary consequences of the individual's actions. It is from this viewpoint that the book brings to- gether a diverse collection of cognitive scientists, behav- ioural ecologists, immunologists and developmental biologists. It is this very diversity that is one of the book's major assets. It is a great pleasure to open a book on evolutionary ecology and find a chapter on cognitive psychology that is both relevant and readable. The volume is divided into five sections: Psycho- logical and cognitive foundations; Communications; Neural, developmental and genetic processes; Hor- monal processes; and the Social context of behaviour. Most chapters are comprehensive reviews of specific behavioural systems such as navigation or song learn- ing, while others develop new models for studying problems in behavioural evolution. Many chapters present primary data and/or new ideas and methods, so at no time does the reader feel that the author is simply regurgitating past publications. The book gets off to a flying start with a radical chapter by Alan Kamil challenging our current approach to the study of animal intelligence. It is his contention that intelligence is adaptive and thus the evolutionary history of the animal is a crucial com- ponent to an animal's ability to learn. Because of this, animal intelligence must be studied in the field rather than in a box in the laboratory. A view echoed by West, King & Freeberg in their chapter on song evol- ution in the cowbird. Kamil sets out clearly his criti- cisms of the old approach, his proposals for a new 'synthetic' approach to the problem and, thankfully, devotes his final section to research strategies that workers should begin to adopt. It would appear that the days of pigeons, rats and bar-pressing are numbered. Kamil's opinions are reinforced by Leslie Real in chapter 5, where the view is expressed that an animal's cognitive architecture is subject to the forces of natural selection just like any other character, a point rather neglected in the past. This opening section on cognition also includes chapters on optimal for- aging as viewed from a learning perspective (Krebs & Inman), navigation in the Hymenoptera (Dyer), and closes with a mathematical chapter illustrating how learning can influence evolutionary change in behav- iour (Papaj). Although this chapter is mathematically quite complex and not for the fainthearted, it is worth persevering because Papaj does eventually dem- onstrate the relevance of the models by application to a real-life bumblebee. With 16 different contributions there are too many to mention each individually in a short review such as this. Suffice to say that not one chapter fails to present high quality research that leaves the reader with much food for thought. It is certainly not a lightweight vol- internal physiological and psychological processes in conjunction with the ecological and evolutionary consequences of the individual's actions. It is from this viewpoint that the book brings to- gether a diverse collection of cognitive scientists, behav- ioural ecologists, immunologists and developmental biologists. It is this very diversity that is one of the book's major assets. It is a great pleasure to open a book on evolutionary ecology and find a chapter on cognitive psychology that is both relevant and readable. The volume is divided into five sections: Psycho- logical and cognitive foundations; Communications; Neural, developmental and genetic processes; Hor- monal processes; and the Social context of behaviour. Most chapters are comprehensive reviews of specific behavioural systems such as navigation or song learn- ing, while others develop new models for studying problems in behavioural evolution. Many chapters present primary data and/or new ideas and methods, so at no time does the reader feel that the author is simply regurgitating past publications. The book gets off to a flying start with a radical chapter by Alan Kamil challenging our current approach to the study of animal intelligence. It is his contention that intelligence is adaptive and thus the evolutionary history of the animal is a crucial com- ponent to an animal's ability to learn. Because of this, animal intelligence must be studied in the field rather than in a box in the laboratory. A view echoed by West, King & Freeberg in their chapter on song evol- ution in the cowbird. Kamil sets out clearly his criti- cisms of the old approach, his proposals for a new 'synthetic' approach to the problem and, thankfully, devotes his final section to research strategies that workers should begin to adopt. It would appear that the days of pigeons, rats and bar-pressing are numbered. Kamil's opinions are reinforced by Leslie Real in chapter 5, where the view is expressed that an animal's cognitive architecture is subject to the forces of natural selection just like any other character, a point rather neglected in the past. This opening section on cognition also includes chapters on optimal for- aging as viewed from a learning perspective (Krebs & Inman), navigation in the Hymenoptera (Dyer), and closes with a mathematical chapter illustrating how learning can influence evolutionary change in behav- iour (Papaj). Although this chapter is mathematically quite complex and not for the fainthearted, it is worth persevering because Papaj does eventually dem- onstrate the relevance of the models by application to a real-life bumblebee. With 16 different contributions there are too many to mention each individually in a short review such as this. Suffice to say that not one chapter fails to present high quality research that leaves the reader with much food for thought. It is certainly not a lightweight vol- ume and undergraduates would, I think, struggle. However, graduate students and experienced researchers alike will, I am sure, benefit from the book. ume and undergraduates would, I think, struggle. However, graduate students and experienced researchers alike will, I am sure, benefit from the book. 790 Book Reviews 790 Book Reviews ? 1995 British Ecological Society, Journal of Animal Ecology, 64, 787-794 ? 1995 British Ecological Society, Journal of Animal Ecology, 64, 787-794 This content downloaded from 169.229.32.136 on Wed, 7 May 2014 17:53:07 PMAll use subject to JSTOR Terms and Conditionshttp://www.jstor.org/page/info/about/policies/terms.jspAll credit to the editor for bringing together researchers from such diverse disciplines of biology studying such a wide range of organisms (slime moulds to primates) in one coherent text illustrating the mechanistic approach to the study of behavioural ecology. RACHEL NEEMS Frank H. Rigler & Robert H. Peters (1995) Science and Limnology. Pp. 239. Excellence in Ecology 6. Ecology Institute, Nordbiinte. DM 74. ISBN 0-932-2205. In the spirit of the Excellence in Ecology Series, Sci- ence and Limnology is a book of advocacy, passion and personal revelation. I did not mind the advocacy but, despite being told in the Preface that the first person singular used throughout the work is a fiction, the personal perspective often gave me a sense of intruding as a reader, which cannot be the intent of a publication! The book was written by Robert Peters but based on the notes of the late Frank Rigler. The main message is that ecology, and ecologists, have lost their way, and nowhere more than in limnology. We are asking the wrong questions at the wrong level, and being less than critical in attempting to answer them. The main problem is that we are being too reductionist in our attempts to come to terms with ecological systems, which throws up com- plexity; what we ought to be doing is focusing on ecological systems as wholes-holism by using sys- tems theory, since this should avoid the complexity, and focus our attention on the more important issues. The holistic approach asserts that: 'Complex systems must be treated as whole systems, because the analysis inherent in reductionism destroys the basis of their integrity'. The idea seems to be to use systems analy- sis-actually correlation and regression analyses in the example presented by the author to illustrate the process in empirical limnology-to expose regularities at the whole-system level that can be used as a basis for prediction and especially for the solution of applied problems. I have at least four problems with this approach. All credit to the editor for bringing together researchers from such diverse disciplines of biology studying such a wide range of organisms (slime moulds to primates) in one coherent text illustrating the mechanistic approach to the study of behavioural ecology. RACHEL NEEMS Frank H. Rigler & Robert H. Peters (1995) Science and Limnology. Pp. 239. Excellence in Ecology 6. Ecology Institute, Nordbiinte. DM 74. ISBN 0-932-2205. In the spirit of the Excellence in Ecology Series, Sci- ence and Limnology is a book of advocacy, passion and personal revelation. I did not mind the advocacy but, despite being told in the Preface that the first person singular used throughout the work is a fiction, the personal perspective often gave me a sense of intruding as a reader, which cannot be the intent of a publication! The book was written by Robert Peters but based on the notes of the late Frank Rigler. The main message is that ecology, and ecologists, have lost their way, and nowhere more than in limnology. We are asking the wrong questions at the wrong level, and being less than critical in attempting to answer them. The main problem is that we are being too reductionist in our attempts to come to terms with ecological systems, which throws up com- plexity; what we ought to be doing is focusing on ecological systems as wholes-holism by using sys- tems theory, since this should avoid the complexity, and focus our attention on the more important issues. The holistic approach asserts that: 'Complex systems must be treated as whole systems, because the analysis inherent in reductionism destroys the basis of their integrity'. The idea seems to be to use systems analy- sis-actually correlation and regression analyses in the example presented by the author to illustrate the process in empirical limnology-to expose regularities at the whole-system level that can be used as a basis for prediction and especially for the solution of applied problems. I have at least four problems with this approach. 1. The fallacy of induction: having I think dismissed induction (making generalizations from cataloguing observations) as the way of achieving scientific under- standing, Peters wants to use correlation and regression-both inductive techniques-as at least foundations for what we do as ecologists. The problem with induction is that no matter how many obser- vations are made on a relationship (swans being white) there is no reason in logic for expecting that the next observation will necessarily conform with the gen- eralization (black swans do turn up); in other words, one should be wary about extrapolating beyond the dataset upon which the correlation/regression analysis was based. So as a foundation for prediction this 1. The fallacy of induction: having I think dismissed induction (making generalizations from cataloguing observations) as the way of achieving scientific under- standing, Peters wants to use correlation and regression-both inductive techniques-as at least foundations for what we do as ecologists. The problem with induction is that no matter how many obser- vations are made on a relationship (swans being white) there is no reason in logic for expecting that the next observation will necessarily conform with the gen- eralization (black swans do turn up); in other words, one should be wary about extrapolating beyond the dataset upon which the correlation/regression analysis was based. So as a foundation for prediction this approach is at best weak. It is strengthened by explor- ing and exposing the mechanistic underpinning of the relationship using the hypothetico-experimental approach. But this is reductionistic. In fairness that is what Peters intends; he wants to use this approach as a basis for creating ideas-hypotheses. But equally in fairness, I do not believe he starts using his holistic approach with a tabula rasa. In the example he gives in his chapter on empirical limnology he seeks a relationship between chlorophyll concentration in lakes and phosphorus concentration. This is surely founded upon preconceived ideas about the relation- ship between the two-mechanisms-which I do not mind, but they are reductionistic. The serious point is that the systems approach advocated might be a start- ing point for science, but it cannot lead in itself to the development of understanding and hence is not a safe basis for making predictions. 2. A fundamental asymmetry in systems analysis: there is a fundamental theorem of systems theory that states that whereas a specific system will lead to a specific relationship between output and input (depen- dent and independent variables), a specific relation- ship between output and input can, in principle, be generated by a whole range of systems. This is the problem of using black box analysis-which is what I think Peters is advocating-to develop a predictive theory (see my number 1). Another way of putting this is that correlations do not prove specific cause- effect relationships! Again, in carrying out the systems analysis and interpreting the results we need to have some ideas about mechanisms-hypotheses-that can be tested. 3. Holism implies integrity in the whole: this is actu- ally written into the definition of holism quoted from the book (above). Yet I am not necessarily convinced of this integrity as far as ecosystems/communities go! They certainly do not have the same integrity as organisms. To imply they do raises all kinds of spec- tres which I am sure Peters does not intend; holism and the organismic approach are, nevertheless, so closely associated that, at the very least, care is certainly needed. There is an important question here, though, that I think Peters misses because of his concerns about reductionism: To what extent do component species-populations contribute to the properties and processes of ecosystems? This is a question that can be, and indeed is being, addressed experimentally by considering predictions from competing models on these relationships (see work reviewed by Fenchel above). 4. Gaps between levels are dangerous: 'reductionism destroys the basis of [the] integrity' of wholes (see above). One possible implication of this is that there are emergent properties (something that I think is not considered by Peters, but I confess that in the absence approach is at best weak. It is strengthened by explor- ing and exposing the mechanistic underpinning of the relationship using the hypothetico-experimental approach. But this is reductionistic. In fairness that is what Peters intends; he wants to use this approach as a basis for creating ideas-hypotheses. But equally in fairness, I do not believe he starts using his holistic approach with a tabula rasa. In the example he gives in his chapter on empirical limnology he seeks a relationship between chlorophyll concentration in lakes and phosphorus concentration. This is surely founded upon preconceived ideas about the relation- ship between the two-mechanisms-which I do not mind, but they are reductionistic. The serious point is that the systems approach advocated might be a start- ing point for science, but it cannot lead in itself to the development of understanding and hence is not a safe basis for making predictions. 2. A fundamental asymmetry in systems analysis: there is a fundamental theorem of systems theory that states that whereas a specific system will lead to a specific relationship between output and input (depen- dent and independent variables), a specific relation- ship between output and input can, in principle, be generated by a whole range of systems. This is the problem of using black box analysis-which is what I think Peters is advocating-to develop a predictive theory (see my number 1). Another way of putting this is that correlations do not prove specific cause- effect relationships! Again, in carrying out the systems analysis and interpreting the results we need to have some ideas about mechanisms-hypotheses-that can be tested. 3. Holism implies integrity in the whole: this is actu- ally written into the definition of holism quoted from the book (above). Yet I am not necessarily convinced of this integrity as far as ecosystems/communities go! They certainly do not have the same integrity as organisms. To imply they do raises all kinds of spec- tres which I am sure Peters does not intend; holism and the organismic approach are, nevertheless, so closely associated that, at the very least, care is certainly needed. There is an important question here, though, that I think Peters misses because of his concerns about reductionism: To what extent do component species-populations contribute to the properties and processes of ecosystems? This is a question that can be, and indeed is being, addressed experimentally by considering predictions from competing models on these relationships (see work reviewed by Fenchel above). 4. Gaps between levels are dangerous: 'reductionism destroys the basis of [the] integrity' of wholes (see above). One possible implication of this is that there are emergent properties (something that I think is not considered by Peters, but I confess that in the absence of an index-the book regrettably does not have one-I cannot be sure). And one interpretation of emergence is that there are gaps between levels of of an index-the book regrettably does not have one-I cannot be sure). And one interpretation of emergence is that there are gaps between levels of 791 Book Reviews 791 Book Reviews ? 1995 British Ecological Society, Journal of Animal Ecology, 64, 787-794 ? 1995 British Ecological Society, Journal of Animal Ecology, 64, 787-794 This content downloaded from 169.229.32.136 on Wed, 7 May 2014 17:53:07 PMAll use subject to JSTOR Terms and Conditionshttp://www.jstor.org/page/info/about/policies/terms.jspArticle Contentsp.790p.791Issue Table of ContentsJournal of Animal Ecology, Vol. 64, No. 6 (Nov., 1995), pp. 675-798+i-viiiVolume Information [pp.795-viii]Front MatterPresidential AddressThe Contribution of Some Recent Research on Birds to Ecological Understanding [pp.675-696]Leaf-Mining Insects and Fluctuating Asymmetry in Elm Ulmus glabra Leaves [pp.697-707]Structure of the Parasitoid Communities of Grass-Feeding Chalcid Wasps [pp.708-720]Differential Response by Males and Females to Brood Manipulations in the Pied Flycatcher: Energy Expenditure and Nestling Diet [pp.721-732]Individual Movements and Estimation of Population Size in Darkling Beetles (Coleoptera: Tenebrionidae) [pp.733-746]A Field Test of Ideal Free Distribution in Flock-Feeding Common Cranes [pp.747-757]Competition for Food in Swans: An Experimental Test of the Truncated Phenotype Distribution [pp.758-766]Synchrony in Tetraonid Population Dynamics [pp.767-776]ForumNo Evidence for Kin-Preferential Swarming in a Daphnia magna Population Coexisting with Fish [pp.777-779]Swarming and Kin in Daphnia: Some Theoretical Considerations [p.780]Identifying Density-Dependent Processes: A Comment on the Regulation of Winter Moth [pp.781-784]Response to Bonsall & Hassell `Identifying Density-Dependent Processes: A Comment on the Regulation of Winter Moth' [pp.785-786]Book Reviewsuntitled [p.787]untitled [pp.787-788]untitled [pp.788-789]untitled [pp.789-790]untitled [pp.790-791]untitled [pp.791-792]untitled [pp.792-793]untitled [p.793]untitled [pp.793-794]Short Notice [p.794]Back Matter [pp.794-iv]