daniel l. alkon, joseph farley,editors, ,primary neural substrates of learning and behavioural...

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Book Reviews 1399 activity budgets of these birds have evolved to minimize energy expenditure, rather than maxi- mize energy intake. Section two deals with the 'Feeding ecology of pelagic marine birds'; the species coverage is rather restricted with three of the seven papers being on sooty shearwaters and two on rhinoceros auklets. Some interesting data on the potential use of olfactory cues by foraging seabirds, a capacity with which they are rarely credited, are presented. The paper by Briggs et al., being a 'joint shil>aircraft- satellite study' of the relationship of phalarope feeding to upwellings off the California coast, demonstrates the way to study seabirds at sea. For most seabird-at-sea workers however, such an approach is prohibitively expensive. As with sec- tion 1, a great deal of information is presented here, and I found the paper by Chu, which examines diet, physiological condition and energy demands in non-breeding sooty shearwaters, particularly inter- esting. On reading section 3, 'Seabird--commercial fisheries interactions', I had a strong, and not wholly inexplicable, feeling of d~jfi vu in places. Many people seem to be worried about potential competition for fish between seabirds and people, but no clear picture emerges from the data on changes in seabird numbers in relation to changes in fisheries. This seems to be due both to difficulties in assessing the availability of fish to seabirds, and their varying capacities to switch prey species when a commonly used resource becomes scarce. A rather dangerous tendency to circularity emerges in the suggestion that, when seabird numbers do not decline in the way expected from fisheries changes, this means that the prey are in fact locally abun- dant. The 10 papers in this section essentially highlight the need for more quantitative data on seabird time budgets and the energetic costs of different activities, since many of the relationships suggested are largely speculative. Data are also presented on seabird mortality in gill nets, much of which could apparently be avoided if, in sensitive areas, such fishing were restricted to the daylight hours. An understanding of the factors affecting prey choice in seabirds suffers from a lack of data on the behaviour and population structures of their prey. Only one paper in this volume is devoted to the dynamics of prey populations, and more communi- cation between fisheries and seabird biologists would clearly be a good move. More than one author suggests that monitoring seabird diets could in fact provide an indication of the demographic structure and distribution of fish populations. We are clearly a long way from this level of understand- ing of the ecological relationships between seabirds and their prey. Still, 'nitimur in adversum' as we say. PAT MONAGHAN Department of Zoology, University of Glasgow, Glasgow G12 8QQ, Scotland. Primary Neural Substrates of Learning and Beha- vioural Change. Edited by DANmL L. ALKON & JOSEPH FARLEY.Cambridge: Cambridge Univer- sity Press (1985). Pp. xi+385. Price s One of the ultimate goals of neurobiology must be the synthesis of explanations of phenomena stu- died and described in terms of behaviour, physio- logy and biochemistry. Synthetic work is hard, because it requires a basic sympathy with the languages and goals of the different brain and behaviour disciplines as they have developed in their own, only partly overlapping modes. It has to avoid premature reduction, the collapse of one language into another, and it demands patience, theoretical and experimental ingenuity to a formid- able degree. Yet few of us would be in this research area if we did not want the chance to accept the synthetic challenge. Of all neural activities in which to search for a likely Rosetta stone, enabling one to read off a translation between the language of behaviour and the language of the cells, learning and memory seem the most promising. When organisms learn, measurable changes occur simul- taneously in their behaviour and their physiology. Surely therefore, we can map one on to the other? Alkon & Farley's book contains a series of reviews summarizing the state of the mapping art, as seen primarily from the Eastern U.S. seaboard. First come the psychologists; Gormezano and Rescorla redefine the Pavlovian laws of learnin~ or rather of conditioning paradigms--within the modern orthodoxy. Garcia, as always iconoclastic and refreshingly literate, offers to liberate us and our research from this conceptual straightjacket. Next, the vertebrate physiologists offer a series of models in which the behavioural terms CS, US and CR are translated into neural pathways and changes in electrophysiological recordings; notable here are the use of the nictitating membrane reflex by Thompson, and of visually conditioned heart rate in the pigeon by Cohen. But the core of the book is, as might be expected from its editors, devoted to the invertebrates and particularly, after a quick and fascinating nod to learning in bees, to the molluscs. All sorts of learning can be demonstrated, it is now clear, in all

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Page 1: Daniel L. Alkon, Joseph Farley,Editors, ,Primary Neural Substrates of Learning and Behavioural Change (1985) Cambridge University Press,Cambridge xi

Book Reviews 1399

activity budgets of these birds have evolved to minimize energy expenditure, rather than maxi- mize energy intake.

Section two deals with the 'Feeding ecology of pelagic marine birds'; the species coverage is rather restricted with three of the seven papers being on sooty shearwaters and two on rhinoceros auklets. Some interesting data on the potential use of olfactory cues by foraging seabirds, a capacity with which they are rarely credited, are presented. The paper by Briggs et al., being a 'joint shil>aircraft- satellite study' of the relationship of phalarope feeding to upwellings off the California coast, demonstrates the way to study seabirds at sea. For most seabird-at-sea workers however, such an approach is prohibitively expensive. As with sec- tion 1, a great deal of information is presented here, and I found the paper by Chu, which examines diet, physiological condition and energy demands in non-breeding sooty shearwaters, particularly inter- esting.

On reading section 3, 'Seabird--commercial fisheries interactions', I had a strong, and not wholly inexplicable, feeling of d~jfi vu in places. Many people seem to be worried about potential competition for fish between seabirds and people, but no clear picture emerges from the data on changes in seabird numbers in relation to changes in fisheries. This seems to be due both to difficulties in assessing the availability of fish to seabirds, and their varying capacities to switch prey species when a commonly used resource becomes scarce. A rather dangerous tendency to circularity emerges in the suggestion that, when seabird numbers do not decline in the way expected from fisheries changes, this means that the prey are in fact locally abun- dant. The 10 papers in this section essentially highlight the need for more quantitative data on seabird time budgets and the energetic costs of different activities, since many of the relationships suggested are largely speculative. Data are also presented on seabird mortality in gill nets, much of which could apparently be avoided if, in sensitive areas, such fishing were restricted to the daylight hours.

An understanding of the factors affecting prey choice in seabirds suffers from a lack of data on the behaviour and population structures of their prey. Only one paper in this volume is devoted to the dynamics of prey populations, and more communi- cation between fisheries and seabird biologists would clearly be a good move. More than one author suggests that monitoring seabird diets could in fact provide an indication of the demographic structure and distribution of fish populations. We are clearly a long way from this level of understand- ing of the ecological relationships between seabirds

and their prey. Still, 'nitimur in adversum' as we say.

PAT MONAGHAN Department of Zoology, University of Glasgow, Glasgow G12 8QQ, Scotland.

Primary Neural Substrates of Learning and Beha- vioural Change. Edited by DANmL L. ALKON & JOSEPH FARLEY. Cambridge: Cambridge Univer- sity Press (1985). Pp. xi+385. Price s

One of the ultimate goals of neurobiology must be the synthesis of explanations of phenomena stu- died and described in terms of behaviour, physio- logy and biochemistry. Synthetic work is hard, because it requires a basic sympathy with the languages and goals of the different brain and behaviour disciplines as they have developed in their own, only partly overlapping modes. It has to avoid premature reduction, the collapse of one language into another, and it demands patience, theoretical and experimental ingenuity to a formid- able degree. Yet few of us would be in this research area if we did not want the chance to accept the synthetic challenge. Of all neural activities in which to search for a likely Rosetta stone, enabling one to read off a translation between the language of behaviour and the language of the cells, learning and memory seem the most promising. When organisms learn, measurable changes occur simul- taneously in their behaviour and their physiology. Surely therefore, we can map one on to the other? Alkon & Farley's book contains a series of reviews summarizing the state of the mapping art, as seen primarily from the Eastern U.S. seaboard.

First come the psychologists; Gormezano and Rescorla redefine the Pavlovian laws of l e a r n i n ~ or rather of conditioning paradigms--within the modern orthodoxy. Garcia, as always iconoclastic and refreshingly literate, offers to liberate us and our research from this conceptual straightjacket. Next, the vertebrate physiologists offer a series of models in which the behavioural terms CS, US and CR are translated into neural pathways and changes in electrophysiological recordings; notable here are the use of the nictitating membrane reflex by Thompson, and of visually conditioned heart rate in the pigeon by Cohen.

But the core of the book is, as might be expected from its editors, devoted to the invertebrates and particularly, after a quick and fascinating nod to learning in bees, to the molluscs. All sorts of learning can be demonstrated, it is now clear, in all

Page 2: Daniel L. Alkon, Joseph Farley,Editors, ,Primary Neural Substrates of Learning and Behavioural Change (1985) Cambridge University Press,Cambridge xi

1400 Animal Behaviour, 33, 4

sorts of molluscans, but the two model systems that dominate the field are Aplysia, in the hands of Kandel, Carew and their collaborators, and Hermissenda from the Woods Hole group headed by Alkon. A substantial portion of the book is taken up by a series of review papers fi'om the two groups on the training paradigms, the physiology, the cells whose plastic properties are being studied, and, finally, some of the biochemistry involved.

For three years or more now the fashionable molecules in the learning field have been cyclic AMP, phosphoproteins, phosphokinases and Ca 2 ions, and in various formulations these are shown to affect or be affected by training procedures that alter ion currents in identified groups of cells. A few years ago it would all have been glycoproteins or neuromodulators. Not represented in this book, but coming up fast, is glutamate. Fashions in biochemicals in learning and memory change as rapidly as they do in depression and schizophrenia.

There are a lot of interesting findings reported in this book, and if some productive model systems in which learning, memory and plasticity studied elsewhere--from the chick to the hippocampal slice--receive no mention, it is still a useful set of papers. But we still wait for a full translation enabling us to identify the necessary, sufficient and exclusive cellular events that 'translate' into a gill withdrawal reflex in Aplysia or association of an odour with a food reward by a bee. Then and only then will it become possible to ask, given a specific set of cellular changes, can we predict the corre- spondent behavioural changes that occur? Despite the new found enthusiasm in learning and memory research, we are a long way off that goal yet.

Biology Department, The Open University, Milton Keynes MK7 6AA, U.K.

STEVEN ROSE

Animal Intelligence. Edited by L. WEISKRANTZ. Oxford: Clarendon Press (1985). Pp. vi+223. Price s

Based upon a Royal Society Meeting held in 1984, this book oilers a timely and high quality survey of the 'state of the art'. There is, however, ample scope for frustration for the average ethologist. Weisk- rantz's optimism was that 'two major s t r a n d ~ careful fieldwork plus the logical dissection of laboratory work--have an opportunity to con- verge . . . ' . But despite repeated references to a century of post-Darwinian progress, there is still a striking lack of convergence here and, behind the simple laboratory-field dichotomy, more funda-

mental adherences are rooted. The most funda- mental of these has to do with whether, at one extreme, 'intelligence' is taken to be whatever capacity an animal has to solve problems that are arbitrary with respect to the forces which have shaped its evolution, or whether it is taken to include capacities that are adapted to a subset of problems special to the animal's ecological niche. Several authors simply never pause to suggest what could possibly be the natural counterpart of the task they set for the animal, and thus what functional sense any reaction makes. This of course is not an inevitable concomitant of controlled experimentation or even of laboratory work, as the papers by, for example, Olton on spatial memory, and Menzel on social foraging, testify. Rather there is a difference between the belief expressed by these authors, that one should initiate research with naturalistic functional analysis, and that expressed by, for example MacKintosh, that 'there is little point in considering such issues until we have done a great deal more to establish that there are any (species) differences to talk about'. But how shall we know what are sensible differences to look for? Perhaps the advantages of working back and forth between functional considerations and a causal/ developmental analysis of learning need yet more advertisement by ethologists (e.g. Bateson 1982). Cheney & Seyfarth are excellent ambassadors here. Their technique of field playback of vocalizations is here turned to the experimental investigation of what vervets know about their social versus physi- cal environments. This represents the first serious testing of the hypothesis that primate intelligence has been honed by the problems set by social interaction, and is accompanied by a nice review of this hypothesis which is much less vague than its predecessors.

On the other hand, fieldworkers like Kummer and Goodall, while offering data which even at an anecdotal level can be extremely important, make little sign of even acknowledging the worries of some experimentalists about what mental capaci- ties can safely be inferred from what behaviours, worries which are synthesized in most extreme and cogent form in McPhail's acceptance of the 'Null Hypothesis' that 'there are no (proven) differences among the intellects of non-human vertebrates'.

This is a fundamental issue, and it is a pity that some editorial initiative could not have ensured a final response from McPhail, following a plethora of apparent contradictions in intervening papers (was it an accident that eight out of 15 papers were on primates?). Likewise, since only one of Terra- ces's two major criticisms of the ape language projects--that there is no evidence for true words nor true sentences--was met, explicit responses by