ECOL 496/596V:

course description
Optional and required readings for this course (pic: (c) NEFSC Photo Archives)
Course syllabus (pdf file)
another course taught by me: ECOL597S
another graduate course taught by me: ECOL467/567
another graduate course taught by me: ECOL596H
Course list for the Department of Ecology and Evolutionary Biology
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Soc. Ins. Lab

Readings for ECOL496V/596V

There are two required textbooks:

•      Krebs JR, Davies NB (1993 for 3rd ed.). An introduction to behavioural ecology. Blackwell Publishers.

This is the standard textbook of behavioral ecology, i.e. studying behavior in an ecological and evolutionary context. It has chapters on foraging, social behavior, reproduction, etc., as well as on the scientific method in behavior research.
Chapters 1, 2, and 3 are available here as pdfs.

•      Shettleworth, S. J., Cognition, evolution, and behavior. 1998, Oxford University Press

This is a textbook that tries to summarize animal research on various aspects of cognition, such as memory and learning, perception, problem-solving, etc. It is bridging the gap between behavioral ecology and (comparative) psychology.

Note that you will not be required to read the whole book(s) (for required readings and dates check timetable on the course description page); both books are classics, easy to read, and summarize the field comprehensively yet concisely. If you are planning to work in an area covered in the course, you'd do well to read these books anyway.

Original papers and scientific reviews

We will also read various original articles to get the current view on various topics.

Comparative analysis:

Greenspan & Swinderen - Cognitive consonance: complex brain functions in the fruit fly and its relatives - Drosophila is discussed as genetic model for studying cognition; indeed much has been learned about mechanisms and costs of learning from studies on the fly. Also discussed are bees and jumping spiders, and the fact that all of these show behaviors and neural signatures that are similar to those shown by mammals when faced with complex tasks.
Menzel & Giurfa - Cognitive architecture of a mini-brain: the honeybee - very nice review showing the many cognitively complex learning tasks that have been set for honey bees, and which they have solved in spite of their very small brains (compared to those of primates).

Is behavior limited by computation costs?
Laughlin - Energy as a constrain on the coding and processing of sensory information - in this review, the energtic demands of the brain (particularly the actual signal transmission in axons) are reviewed (20% for humans), with a focus on the design of vision circuitry and its optimization to save energy. Particularly interesting: the cost ratio between maintaining neurons and producing signals on them determines whether there should be one neuron for each event ('grandmother neuron') or each event shoudl be identified by a unique combination of neurons.
Mery & Kawecki - A fitness cost of learning ability in Drosophila melanogaster - this is one of a pair of papers by the same authors on costs of learning, both in terms of productions and maintenance of neural tissue, and of actually using it. It had been known that both these things may be costly, but these are the first studies that show this directly and clearly though artificial selection experiments.

Does brain size predict behavioral complexity? (If yes, is it absolute or relative brain size?)
Dunbar - The social brain hypothesis - Robin Dunbar is the major proponent of the hypothesis that large brains evolved to deal with large group sizes (and more complex social interactions). He reviews here what other hypotheses there are, the data for primates, and what measures of brain size may be used. His hypothesis holds in particular for neocortex size/size of rest of brain. The role of body size in the evolution of large brains is discussed, but no major solution found, although he says that brain size may be more conserved than body size.
Marino - Absolute brain size: did we throw the baby out with the bathwater? - current information on the use of relative or absolute brain size in judging cognitive or behavioral complexity. It seems absolute brain size correlates with number of glia (supporting) cells and energy expenditure. Larger brains thus may necessitate more complex architecture - though whether this architecture then contributes to cognition or just to retaining the same functions as smaller brains is unclear.
Mares et al. - Brain allometry in bumblebee and honey bee workers - this is a study looking at variation in brain size within a species (bumble bees) and also comparing it to another species, honey bees. The general trend that larger animals have larger brains is supported here. However, it is not clear whether this predicts higher behavioral complexity in larger animals.
Gronenberg et al. - Age-dependent and task-related morphological changes in the brain and the mushroom bodies of the ant Camponotus floridanus - brain volume, especially of mushroom bodies, can increase in ants depending on experience. This is probably due to axon and dendrite growth rather than cell proliferation. Larger ants don't necessarily have larger brains (within the same species).

Do structural constraints influence behavior? (What do we know about actual neural mechanisms?)
Delcomyn - Walking robots and the central and peripheral control of locomotion in insects - investigating the neural networks necessary to make an insect (or a robot) walk is a major research field, and walking is one of the behaviors that are best understood in terms of the network design necessary to achieve it.
Chittka et al. - Flower constancy, insect psychology, and plant evolution - flower constancy is the behavior of bees that avoid flowers of different species in favor of visiting just one species, while other bees of the same colony specialize on other flower species. This may in part be caused by a limited working memory, and additional costs if the bee wants to 'upload' handling strategies for a different flower type.
Bernays - Neural limitations in phytophagous insects: implications for diet breadth and evolution of host affiliation - classic paper, for the first time discussing neuronal limitations as cause for dietary specialization in insect species. This is NOT flower constancy, Bernays talks about insect species that exclusively or mostly eat one plant species.

Pinker - Why nature and nurture won't go away - an essay discussing the history of the debate and its philosophical and political implications (e.g. on justice and punishment, on racism and sexism, etc.). Pinker clearly states that any human behavioral trait is the result of an interplay of genetic heritage and environmental influence, but at the same time stresses that researchers should continue to disentangle these effects for each particular behavior. In addition, he points out, many scientists still promote a sort of hidden 'only nurture' paradigm: for example, Stephen Jay Gould saying that the human “brain [is] capable of a full range of behaviors and predisposed to none.”
Robinson et al. - Sociogenomics: social life in molecular terms - a review of which genes are known to influence behavior, particularly social behavior (all examples in a summary table are either from insects or mice/rats, although some microbes are also discussed). This article gives a wealth of interesting examples, and in many cases of genes that are shown to have similar effects across taxa. It also discusses that looking at transcription (i.e. quantifying RNA products or particular genes) is not sufficient to understand heritability, because environmental or behavioral changes can cause changes in transcription rates (rather than the other way around), making the causal link uncertain. However transcription databases certainly are helpful in investigating which genes/enzymes may be involved in the physiological processes that underlie behaviors.
The Honeybee Genome Sequencing Consortium - Insights into social insects from the genome of the honeybee Apis mellifera - the paper that described the sequencing of the honey bee genome (Apis mellifera, in 2006; est. 10000 genes). This is a landmark paper because it adds the first social insect to the set of genetic 'model organisms'. Some important differences in the organisation of this genome compared to Drosophila, the fruit fly, are found - but also many similarities. The paper also describes some conclusions for evolution of the subspecies of A. mellifera.
Brakefield - Evo-devo and constraints on selection

Game theory:
Nowak & Sigmund - Evolutionary dynamics of biological games - a review on the current uses of game theory in biology. This is an excellent short summary of classical game theory approaches, but its strength is particularly in detailing new approaches, such as the analysis of replicator and adaptive dynamics. The latter involves studying the states that a population is likely to evolve toward in a continuous, rather than small and discrete, strategy space.
Barraclough et al. - prefrontal cortex and decision making in a mixed strategy game
Lewis - Beyond dominance: the importance of leverage

Predation and parasitism:

Dall et al. - Information and its use by animals in evolutionary ecology

Behavioral syndromes:
Sih et al. - Behavioral syndromes: an ecological and evolutionary overview - the main author of this paper is one of the prime proponents in studying 'behavioral syndromes', something like personalities in animals, and he indeed coined this term. The review explains terms and gives many examples.
Figueredo et al. - Consilience and life history theory: from genes to brain to reproductive strategy - a somewhat long paper, but with the interesting thesis that the many identified human 'personality' dimensions can maybe be unified into an r-k dimension: a single number that reflects how much towards an r- or K- (in the ecological sense of reproducing early and a lot with little investment - r - or the opposite - K) strategist a person behaves. The authors show that their aggregate parameter does predict a lot of behaviors in mating, reproduction, health, and other contexts.
Bell - Future directions in behavioural syndromes research

Evolution of cooperation & altruism:
Linksvayer & Wade - The evolutionary origin and elaboration of sociality in the aculeate hymenoptera: maternal effects, sib-social effects, and heterochrony - why did bees evolve sterile workers and cooperation in general? Are they predisposed to this because of their haplodiploidy, or are workers the result of parental manipulation? A comprehensive review.
Fehr & Rockenbach - Human altruism: economic, neural, and evolutionary perspectives - why are humans so much more altruistic than animals, and than predicted by conventional economic theories? The experiments on this phenomenon, as well as possible explanations, are reviewed. See also the other paper by the same first author below.
Komdeur - Variation in individual investment strategies among social animals

Living in groups:
Walker et al. - Does soil type drive social organization in southern hairy-nosed wombats? - Wombats! Hairy-nosed! How can you go wrong with this. A paper investigating the environmental factors that can lead to sociality in animals.

Krebs & Dawkins - Mind-reading or manipulation - in this chapter from an edited textbook the authors put up the hypothesis that (most) animal communication is not 'designed' to maximise information transfer, but that instead signal senders want to manipulate the receiver, whereas receivers only pay attention to signals in order to predict what the sender is going to do next ('mind-reading'). Classic!
Zahavi - Mate selection - selection for a handicap - the original article (cited more than 1000 times) in which Zahavi developed his handicap theory of animal signals. Essentially this states that a signal has to be costly to ensure reliability (honesty). This has primarily been used to explain extravagance in sexual signals, but applies to other kinds of signalling as well.
Hurd & Enquist - A streategic taxonomy of biological communication

Human behavioral ecology:
Aiello & Dunbar - Evolution of language - can what we know about brain size in primates help us predict what group size humans lived in early in their evolution? And what are the implications for language evolution in humans?
Fehr & Fischbacher - The nature of human altruism - why are humans so much more altruistic than animals, and than predicted by conventional economic theories? The experiments on this phenomenon, as well as possible explanations, are reviewed.
Bird et al. - The hunting handicap: costly signaling in human foraging strategies - applying animal signaling theory to (contemporary) human societies.

The next questions in behavioral ecology:
Read & Clark - The past 20 years of ecology and evolution - one-page look back at the world 20 years ago, and how much more relevant challenges the study of ecology faces today.
Read & Clark - The next 20 years of ecology and evolution - again, some current issues (habitat destruction, climate change, antibiotic resistance, etc.) are mentioned. Other trends are identified: systems biology, immunology, biomedicine, the tension between model organisms and studying their ecology and natural history, evolution of sex. Finally, it advocates more communication with the public about scientific methods and results.
Owens - Where is behavioural ecology going? - nice review of both traditional and new approaches and questions in behavioral ecology. Main point is that behavioral ecologists are likely to (or at least would benefit from) increased study of mechanisms, i.e. gene-behavior mappings, and use of model organisms. The lack of replication of studies on diverse species is stated as a main failing of the field.
Clark & Gelfand - the authors argue that new modeling approaches should integrate information from more sources and more effects. Mostly, a particular approach, Bayesian inference is advocated.

Cognition Introduction
Pennisi - Social animals prove their smarts - I highly recommend this short article on the history and currently active areas of the study of animal cognition. The article somewhat focuses on sociality promoting cognitive abilities, but not only this is discussed.

Perception & attention
Dukas - Causes and consequences of limited attention - a review of the author's and some other studies on (mostly jay) perception and attention. Attention here is described as a limited part of the visual field that can be inspected at any given time, with a resolution that is dependent on how much brain tissue is devoted to this task. The author then presents a model on how the amount of brain tissue and the visual angle used may have evolved to maximize predator detection in a limited amount of time.
Spaethe et al. - Do honeybees detect color targets using serial or parallel visual search? - Visual search refers to the problem of finding a visual 'target' among 'distractors'. In humans and some other mammals, when the target differs along only one dimension (e.g., color, not shape), the detection time is independent of number of distractors ('parallel search' - because all objsects seem to be scanned in parallel). When two dimensions are involved, search is serial (objects appear to be scanned in sequence, and decision time is proportional to number of distractors). Here, Spaethe et al. find that bees seem to use only serial search, even when only color differs between objects. They also show that detection is severely impeded by distractors even when the color contrast between targets and distractors is significant. (Note the use of the color hexagon, a way of plotting the response of the three bee color receptors for any color.)
Tautz et al. - Honeybee odometry: performance in varying natural terrain - Earlier studies had shown that bees measure flown distance by measuring optic flow (using only the green receptor - this is also reviewed here). This study shows the implications: that bee distance estimates are highly context dependent, but that inter-bee communication still works because they all use the same method for distance estimation. This issue is an example for how mechanism-dependent understanding of perception and resulting behavior can be - one cannot understand the waggle dance and bee orientation behavior over water unless one takes the perception mechanism into account.
Spaethe & Chittka - Interindividual variation of eye optics and single object resolution in bumblebees - Visual acuity in bees depends on body size as well as peripheral processing mechanisms, such as neuronal pooling between retina and higher-level information processing centers. In this case, larger bumble bees can detect smaller targets (or target from a larger distance) than small ones, with implications for division of labor as well as the evolution of body size.
Chittka et al. - Flower constancy, insect psychology, and plant evolution - (also discussed above in the Neurobiology section) The concept of attention has a lot of similarity with discussions of limited working memory. This paper explores some ecological consequences of such limitations of immediate information processing.
Bernays - Neural limitations in phytophagous insects: implications for diet breadth and evolution of host affiliation - (also discussed above in the Neurobiology section) Like the previous paper, this paper deals with ecological and evolutionary consequences of limitations in processing of sensory information. Although this may not be said explicitly, the idea that insect herbivores may specialize to avoid costly neural flexibility is essentially the same as what Dukas says about attention: 'attentional focus' may evolve to save brain tissue costs.
Greenspan & Swinderen - Cognitive consonance: complex brain functions in the fruit fly and its relatives - (also discussed above in the Neurobiology section) Page 3-4 give a discussion of new results on possible neuronal signatures of 'attention' in flies as well as reviewing those from mammals.

Learning and memory
Menzel & Giurfa - Cognitive architecture of a mini-brain: the honeybee - stages of memory and neuronal correlates (this has been studied most extensively in honey bees).
Greenspan & Swinderen - Cognitive consonance: complex brain functions in the fruit fly and its relatives - (already mentioned twice!) Check the first part of this paper for a brief discussion of declarative memory (here called explicit memory): cognitive maps in honey bees and anticipatory learning in spiders.
Cheng - A purely geometric module in the rat's spatial representation - Is the mind composed of modules working in parallel on specific tasks? The geometric module is one of the main pieces of evidence that this may be true in some contexts. See the Shettleworth book, chapter 3, for a brief summary of this study (Fig 3.2, see also fig. 7.9).
Clayton & Krebs - Memory for spatial and object-specific cues in food-storing and non-storing birds - Is learning ability species-specific and task-specific? This study provides evidence for both, and thus for the hypothesis that there is no general 'learning module', but instead animals are adapted to solve particular tasks relevant to them using learning.

Orientation & Navigation
Menzel et al. - Two spatial memories for honeybee navigation - Do honeybees have cognitive maps? This was a long discussion (in which Menzel has argued that there is not sufficient evidence that they do). However, with this new paper, it seems that bees do have an allocentric, general landscape memory (a cognitive map?); they just don't use it if they have been extensively trained to one location, in which case they use their 'autopilot' of vector memory.

Classification & generalization
Greenspan & Swinderen - Cognitive consonance: complex brain functions in the fruit fly and its relatives - Check pages 2-3 for a brief review of categorization and abstract concepts in honey bee visual learning tests.

Free will?
Glimcher - Indeterminacy in brain and behavior

Other books on animal behavior worth reading (biased list by me)

Introductory books on social behavior.

•      Gadagkar R (1997) Survival strategies: cooperation and conflict in animal societies. Publisher: Harvard University Press.

Brief, but comprehensive and easy to read introduction to social behavior. Lots of great and original examples of cooperation and conflict particularly, but not only, from social insects.

Books on methods

•      Martin P, Bateson P (1993) Measuring behaviour. Cambridge University Press.

Different methods of quantifying behavior and designing experiments.

•      Dytham C (2003) Choosing and using statistics: a biologist’s guide. Blackwell Publishing.

One of the many books that try to explain stats to biologists. You will have to read a stats book eventually if you haven’t already done it. Pay attention to the general insights, like types of data, types of tests, and how to find out which test to use.

Introductory and more specialized books on social insects.

•      Hölldobler B, Wilson EO (1995). Journey to the ants: a story of scientific exploration. Publisher: Belknap Press.

Lovely book written by ant-enthusiasts about the spectacular and strange behaviors shown by ants. Easy to read. A must for those who want to know what little social insects can achieve with collective action.

•      Heinrich, B (1979) Bumblebee economics. Harvard University Press has reprints from 2004.

Best introduction to bumble bees. Starts with general natural history, then talks about thermoregulation, foraging, development, etc.

•      Wilson EO (1974) The insect societies. Belknap Press.

Old but comprehensive and nicely illustrated introduction to social insects. If you don’t know social insects, you can find out about all the different groups and their most interesting representatives here.

•      Hölldobler B, Wilson EO (1990) The ants. Belknap Press.

The ant-bible. Very detailed, review-like, but not totally up-to-date. Nevertheless, good first resource to find out what is known about a topic. Also has descriptions of all ant groups.

•      Gordon D (2000) Ants at Work: How an Insect Society is Organized. W W Norton & Co.

Little, easy read book on task allocation in harvester ants and how she found out about it.

•      Seeley T (1996) The Wisdom of the Hive: The Social Physiology of Honey Bee Colonies. Harvard University Press.

Excellent comprehensive book on honey bees and how they forage and communicate. Lots of detail on collective behavior in honey bees.

•      Seeley (1985) Honeybee Ecology: A Study of Adaptation in Social Life. Princeton University Press.

•      Frisch Kv (1967) The dance language and orientation of bees – reprinted by Belknap Press 1993.

The classic bee bible. This contains a summary of the first studies on honey bee dances and many aspects of their behavior. Today much seems anecdotal, but even today this is the first resource to look for what we know on honey bee behavior.

•      Goulson D (2003) Bumblebees: their behaviour and ecology. Oxford University Press.

More details, and more up-to-date, on bumble bees.

•      Camazine S, Deneubourg J-L, Franks NR, Sneyd J, Theraulaz G, Bonabeau E (2003). Self-Organization in Biological Systems (Princeton Studies in Complexity). Publisher: Princeton University Press.

This is a fairly detailed and extensive treatment of the topic of self-organization, i.e. the emergence of collective behavior without a central leader or template. Most examples are from social insects, but there are also some from microbes etc. Written by the world experts in the field which, may mean that it’ll be slow reading for beginners. Nevertheless a great resource.