Towards a field theory of behaviour

Classical ethology, with its emphasis on separability of parts, has largely failed to do justice to the wholeness of the individual animal, to the integrity of group behaviour and to the continuity between observable behaviour and consciousness. Field theory has potentialities to do better, as illustrated in this paper with reference to morphogenetic and behavioural fields. A behavioural domain is delineated — playlike behaviour — where field theory is particularly relevant. It is shown that the concept of symmetry can suggest new questions as well as explain some generally known phenomena of group behaviour. New interpretations of displacement activities and of etho-ecological adaptations are offered, both of which involve the whole individual animal.This paper is dedicated to W.M.S. Russell at the occasion of his retirement. He and Claire Russell invented most of what is new in ethology years or decades ago.     

Zur m?glichen Funktion und Konvergenz wippender K?rperbewegungen bei V?geln

Summary Many species of wagging birds — e.g. tail-wagging Common and Spotted Sandpiper (Actitis hypoleucos, A. macularia), some Wagtails (Motacillidae), Forktails (Enicuridae) or body-bobbing Dippers (Cinclidae) — inhabit fast-running streams and it appears that the behaviour is a convergent adaption to this habitat. Possible functions are discussed; the hypothesis is given that birds of fast-running streams benefit due to wagging as an optical intraspecific signal (e. g. in territorial or sexual behaviour). Owing to the unfavourable acoustical situation (noise of the running water) and the linear habitat, vocal signals are presumed to be generally less effective than optical signals. Predictions concerning the European species based on this hypothesis are in agreement with the available evidence in literature.     

Mapping of a mouse homolog of a Heterochromatin protein gene to the X Chromosome

Modifiers of position-effect-variegation inDrosophila are thought to encode proteins that are either structural components of heterochromatin or enzymes that modify these components. We have recently shown that a sequence motif found in oneDrosophila modifier gene, Heterochromatin protein 1 (HP1), is conserved in a wide variety of animal and plant species (Singh et al. 1991). Using this motif, termed chromo box, we have cloned a mouse candidate modifier gene,M31, that also shows considerable sequence homology toDrosophila HP1. Here we report evidence of at least four independently segregating loci in the mouse homologous to theM31 cDNA. One of these loci—Cbx-rsl—maps to the X Chromosome (Chr), 1 cM proximal toAmg and outside the X-inactivation center region.     

Transfer of a time-signal isochronous with local time in translocation experiments to the geographical longitude

Summary When mushroom-bodies of time-trained honeybees are translocated according to the geographical longitude — the latitude remains nearly the same — and are implanted to naive host animals the information on time is transferred as an information on local time, i.e. the time-signal learned by the donor animal breaks through in the host animal isochronous with local time and not as an endogenously driven circadian rhythm.In memoriam of Prof. Dr. K. v. Frisch on the occasion of the 100th birthday     

Mutants as tools to understand cellular and molecular drought tolerance mechanisms

The use of mutants: a most promising way to detect genes involved in development or in response to environmental stress. The model species Arabidopsis, particularly amenable to dissect the genetics and molecular mechanisms underlying physiological responses, also offers the advantage of a wide variety of mutants. As far as drought tolerance is concerned, hormonal mutants, impaired in hormone biosynthesis — deficient mutants — or in the signal transduction pathway—responsive mutants—provide a valuable tool to analyse the role of phytohormone interaction in the plant drought behaviour as well as to differentiate the mutant phenotypes with new criteria.These two categories of mutants (in particular the abscisic acid, ABA, mutants) were shown to be affected in developmental processes during seed maturation-in the desiccation phase- and/or in response to environmental stress (drought, ...) in vegetative tissues. The present report will focus on this last aspect: alterations in drought responses in vegetative tissues (adaptive strategies and drought tolerance mechanisms) essentially in Arabidopsis hormonal mutants (ABA-deficient and ABA-insensitive, GA-deficient, auxin and ethylene-insensitive).Some of the results are discussed with regard to the predicted functions of genes affected by the mutations.     

Labelling of regenerating retinal pigment epithelium by colloidal iron oxide and ferritin conjugated to wheat germ agglutinin

Summary In order to determine if there are biochemical changes in plasma-membrane oligosaccharides of regenerating retinal pigment epithelium, the binding of colloidal iron oxide at low pH and ferritin-conjugated wheat germ agglutinin — probes of sialic acid and N-acetylglucosamine on the cell surface — was examined electron-microscopically. An animal model of retinal pigment epithelium regeneration — rabbits with sodium iodate induced retinopathy — was used. In this model, large expanses of regenerating pigment epithelium are present for comparison with zones of spared pgiment epithelium in the same animals. In thin sections examined by transmission electron microscopy, ferritin-conjugated wheat germ agglutinin appeared to bind more intensely to the exposed plasma membrane of regenerating retinal pigment epithelium than to spared pigment epithelium, or that of normal rabbits. Morphometry verified this. Colloidal iron oxide intensely labelled the plasma membranes of regenerating, spared, and normal pigment epithelium, and was visibly reduced after exposure of tissue to neuraminidase. The observations indicate that the plasma membrane of regenerating retinal pigment epithelium bears sialic acid and N-acetylglucosamine residues as in normal retinal pigment epithelium. However, the amount of plasma membrane bearing exposed N-acetylglucosamine increases during regeneration.     

Plant cells — young at heart?

Dolly has become a synonym for one of the greatest breakthroughs in animal reproductive biology: the regeneration of a whole mammal from a somatic cell nucleus. The equivalent experiments in plants — the regeneration of whole plants from single differentiated cells — are comparatively easy. Does this apparent difference in the developmental potential of animal and plant somatic cells reflect mechanistic differences in the regulation and maintenance of their respective cell differentiation?     

Individual boldness is linked to protective shell shape in aquatic snails

The existence of consistent individual differences in behaviour (‘animal personality’) has been well documented in recent years. However, how such individual variation in behaviour is maintained over evolutionary time is an ongoing conundrum. A well-studied axis of animal personality is individual variation along a bold–shy continuum, where individuals differ consistently in their propensity to take risks. A predation-risk cost to boldness is often assumed, but also that the reproductive benefits associated with boldness lead to equivalent fitness outcomes between bold and shy individuals over a lifetime. However, an alternative or complementary explanation may be that bold individuals phenotypically compensate for their risky lifestyle to reduce predation costs, for instance by investing in more pronounced morphological defences. Here, we investigate the ‘phenotypic compensation’ hypothesis, i.e. that bold individuals exhibit more pronounced anti-predator defences than shy individuals, by relating shell shape in the aquatic snail Radix balthica to an index of individual boldness. Our analyses find a strong relationship between risk-taking propensity and shell shape in this species, with bolder individuals exhibiting a more defended shell shape than shy individuals. We suggest that this supports the ‘phenotypic compensation’ hypothesis and sheds light on a previously poorly studied mechanism to promote the maintenance of personality variation among animals.     

Attempt to modify rate and duration of licking in rats by operant conditioning

Lick-rate in rats is said to be constant for a given animal, despite variations of internal and external stimuli. On the other hand, small changes can be observed due to changes in the construction of the licking device. However, variations do not exceed 20%.In an attempt to gain operant control over the ILI (interlick interval — the time between two lick-onsets) the delivery of reinforcement (20 μl water) was made dependent on the emission of ILIs of a predetermined length longer than during baseline licking. It could be observed that rats could not shift the peak of their ILI distribution within the reinforced range but — to increase the number of reinforcements — they increased the scatter of the ILI distribution or developed a “harmonic” peak at double ILI length. When the animals were forced in a second experiment to prolong the lick-duration (time of tongue-spout contact) to obtain water, they failed if the restriction from the drinking spout made a closer approach impossible.It is argued that the ability to obtain reinforcement under both schedules is due to postural changes of the animal. The mechanisms controlling licking seem to be relatively constant, which allows good coordination with other behaviours which have to be performed during drinking, such as breathing and swallowing. It can be concluded that the amount of water consumed by rats is controlled by the length of time spent in licking and not by changing the lick-rate.     

The contribution of additive genetic variation to personality variation: heritability of personality

Individual animals frequently exhibit repeatable differences from other members of their population, differences now commonly referred to as ‘animal personality’. Personality differences can arise, for example, from differences in permanent environmental effects―including parental and epigenetic contributors―and the effect of additive genetic variation. Although several studies have evaluated the heritability of behaviour, less is known about general patterns of heritability and additive genetic variation in animal personality. As overall variation in behaviour includes both the among-individual differences that reflect different personalities and temporary environmental effects, it is possible for personality to be largely genetically influenced even when heritability of behaviour per se is quite low. The relative contribution of additive genetic variation to personality variation can be estimated whenever both repeatability and heritability are estimated for the same data. Using published estimates to address this issue, we found that approximately 52% of animal personality variation was attributable to additive genetic variation. Thus, while the heritability of behaviour is often moderate or low, the heritability of personality is much higher. Our results therefore (i) demonstrate that genetic differences are likely to be a major contributor to variation in animal personality and (ii) support the phenotypic gambit: that evolutionary inferences drawn from repeatability estimates may often be justified.     

Editorial

These are heady times. As the new millennium dawns, the field of zoology is more exciting than ever. Phylogenetic relations among animals are rapidly being refined or revised in light of new data and powerful new analyses. These revelations in turn are rekindling interest in the function, development, and evolution of animals from all branches of the zoological tree. Tools and insights derived from work on a few model organisms are being applied widely to fill in significant gaps in our knowledge about how the astonishing diversity of animal characteristics evolved.Paralleling this new excitement is an upheaval in scientific publishing. Increasingly specialized journals are sprouting up worldwide as publishers rush to embrace emerging areas of excitement. Electronic communication is re-defining expectations about how new knowledge is disseminated both in format and in mode of transmission and about the time from submission to publication. For established journals to survive, they too must evolve.This journal, ZOOLOGY, is evolving to meet the needs and expectations of the modern community of animal biologists. It aims to promote research like that of it's founder J.W. Spengel — that explicitly emphasizes comparative aspects of animal biology. With the rapid growth of phylogenetic information, ZOOLOGY now offers a venue where the full impact of recent phylogenetic advances on our understanding of animal form, function, development, and evolution can be addressed. An ambitious new group of editors, a new advisory board, and a professionally staffed editorial office, are working to rejuvenate ZOOLOGY as an internationally recognized leader in comparative animal biology.ZOOLOGY has a distinguished place in the pantheon of animal biology journals. It ranks among the oldest continuously published journals in Germany. Founded in 1886 as Zoologische Jahrbücher it quickly rose to prominence among European zoological journals. In the early 1990's, after political and social upheavaling some European countries and after the reunification of Germany, the journal took a more international approach under a new title, ZOOLOGY — ANALYSIS OF COMPLEX SYSTEMS (ZACS). For the last four years, ZOOLOGY has also worked in close cooperation with the German Zoological Society and this tradition will continue. Once each year, it will publish review lectures presented at the annual meeting of the Society. A supplement to ZOOLOGY, containing the abstracts of oral and poster presentations of the annual meeting, will also be produced in conjunction with the German Zoological Society.But the new ZOOLOGY also recognizes the increasing importance of rapid and international communication in all areas of animal biology. It therefore aims to reduce publication time drastically and to enhance the speed and rigor of the review process. The new editors and editorial board are committed to maintaining the highest scientific standards and also to remaining flexible enough to adjust to the ever changing field of animal biology. Recognizing the increasing importance of rapid, effective, international communication, the new ZOOLOGY will offer:— the highest scientific standards— a short review time— a publication time schedule of three months after acceptance— color plates free of charge (at the editors' discretion)— an online version published in advance of the printed journal— free E-mail Table of Contents alerts so papers are widely publicized— papers abstracted/indexed by all the major scientific indexing servicesAs the new cover and the more accessible layout inside this issues show, the change has already begun. The future holds great promise for animal biology. Join us as Zoology strives to fulfil that promise.Spring 2001J. Matthias StarckThomas C. BoschA. Richard PalmerKiyokazu Agata     

Identifying Social Learning in Animal Populations: A New ‘Option-Bias’ Method

Background

Studies of natural animal populations reveal widespread evidence for the diffusion of novel behaviour patterns, and for intra- and inter-population variation in behaviour. However, claims that these are manifestations of animal ‘culture’ remain controversial because alternative explanations to social learning remain difficult to refute. This inability to identify social learning in social settings has also contributed to the failure to test evolutionary hypotheses concerning the social learning strategies that animals deploy.

Methodology/Principal Findings

We present a solution to this problem, in the form of a new means of identifying social learning in animal populations. The method is based on the well-established premise of social learning research, that - when ecological and genetic differences are accounted for - social learning will generate greater homogeneity in behaviour between animals than expected in its absence. Our procedure compares the observed level of homogeneity to a sampling distribution generated utilizing randomization and other procedures, allowing claims of social learning to be evaluated according to consensual standards. We illustrate the method on data from groups of monkeys provided with novel two-option extractive foraging tasks, demonstrating that social learning can indeed be distinguished from unlearned processes and asocial learning, and revealing that the monkeys only employed social learning for the more difficult tasks. The method is further validated against published datasets and through simulation, and exhibits higher statistical power than conventional inferential statistics.

Conclusions/Significance

The method is potentially a significant technological development, which could prove of considerable value in assessing the validity of claims for culturally transmitted behaviour in animal groups. It will also be of value in enabling investigation of the social learning strategies deployed in captive and natural animal populations.     

CD19 maps to a region of conservation between human chromosome 16 and mouse chromosome 7

CD19 is a B lymphocyte cell surface protein expressed from the earliest stages of B lymphocyte development unitl their terminal differentiation into plasma cells. In this report the human CD19 gene (hCD19) was localized to band p11.2 on the proximal short arm of chromosome 16 by in situ hybridization to metaphase chromosomes, using hCD19 cDNA as probe. hCD19 gene localization was confirmed by polymerase chain reaction based analysis with hCD19-specific primers, using a panel of human/hamster somatic cell hybrid DNA as templates. The mouse CD19 gene (MCd19) was mapped to bands F3-F4 of chromosome 7 by in situ hybridization to metaphase chromosomes, using a mCD19 cDNA probe. Segregation analysis of nucleotide sequence polymorphisms in inter-specific backcross progeny revealed linkage of mCd19 with hemoglobin (Hbb), Int-2, and H19, other loci previously mapped to the same region of mouse chromosome 7, confirming the localization of mCd19 to this region. The order of these loci was determined to be centromere — Hbb — mCd19 — H19 — Int-2 —telomere. The genetic distance between the loci examined, calculated from the recombination frequencies, suggested that mCd19 was located centrally between Hbb and H19. This region of mouse chromosome 7 is homologous to the region of human chromosome 16 to which the hCD19 gene maps. Multiple genes with a lymphocyte-related function also map to this conserved region including genes encoding the IL-4 receptor, CD11a, CD11b, CD11c, CD43 (leukosialin), and protein kinase C polypeptide.     

Impedance traces of copepod appendage movements illustrating sensory feeding behaviour

An experimental system incorporating a computerized micro-impedance unit has been used to make direct measurements of the activity of copepod cephalic appendages. As the appendages are used to both propel the copepod through water and handle particles, it follows that appendage activity reflects feeding behaviour.To investigate the sensory feeding behaviour of copepods, their activity was recorded with food stimuli varying in size and chemical composition. Sample impedance traces are given for the appendage movements of Temora longicornis in the presence of: 1 — filtered seawater; 2 — beads; 3 — phytoplankton cells; 4 — dissolved free amino acids. The normal appendage movements shown in filtered seawater were modified when copepods were offered particles and dissolved chemicals. Results show that chemical and mechanical stimuli are responsible for the recognition and selection of food. Impedance traces distinguish between behavioural responses such as: antennule flicks, leg kicks, combing, handling and rejection of particles. Spectral analyses of traces have demonstrated that differences in beat pattern are significant.Paper presentation, at the Third International Conference on Copepoda. British Museum (Natural History), London, U. K., 10–14 August, 1987.     

Generation time and albumin evolution

The albumins of long-generation animals such as man and chimpanzee are shown to be no less evolved immunologically than those of some short generation time rodents such as rat, mouse, and hamster. This finding casts very serious doubt on certain assumptions concerning the times of divergence of these species made by DNA hybridization workers. As these assumptions have led them to conclude that generation time is a key factor controlling the rate of genomic or DNA evolution, this latter conclusion is also made suspect. The molecular clock concept—that it is absolute time which determines the amount of protein and DNA evolution—is, on the other hand, strengthened.This research was supported by NSF grants GB-20750 to the author and GB-13119 to Dr. A. C. Wilson.     

Keeper-Animal Interactions: Differences between the Behaviour of Zoo Animals Affect Stockmanship

Stockmanship is a term used to describe the management of animals with a good stockperson someone who does this in a in a safe, effective, and low-stress manner for both the stock-keeper and animals involved. Although impacts of unfamiliar zoo visitors on animal behaviour have been extensively studied, the impact of stockmanship i.e familiar zoo keepers is a new area of research; which could reveal significant ramifications for zoo animal behaviour and welfare. It is likely that different relationships are formed dependant on the unique keeper-animal dyad (human-animal interaction, HAI). The aims of this study were to (1) investigate if unique keeper-animal dyads were formed in zoos, (2) determine whether keepers differed in their interactions towards animals regarding their attitude, animal knowledge and experience and (3) explore what factors affect keeper-animal dyads and ultimately influence animal behaviour and welfare. Eight black rhinoceros (Diceros bicornis), eleven Chapman’s zebra (Equus burchellii), and twelve Sulawesi crested black macaques (Macaca nigra) were studied in 6 zoos across the UK and USA. Subtle cues and commands directed by keepers towards animals were identified. The animals latency to respond and the respective behavioural response (cue-response) was recorded per keeper-animal dyad (n = 93). A questionnaire was constructed following a five-point Likert Scale design to record keeper demographic information and assess the job satisfaction of keepers, their attitude towards the animals and their perceived relationship with them. There was a significant difference in the animals’ latency to appropriately respond after cues and commands from different keepers, indicating unique keeper-animal dyads were formed. Stockmanship style was also different between keepers; two main components contributed equally towards this: “attitude towards the animals” and “knowledge and experience of the animals”. In this novel study, data demonstrated unique dyads were formed between keepers and zoo animals, which influenced animal behaviour.     

High-resolution linkage map in the vicinity of the Lp locus

Looptail (Lp) is a mutation that profoundly affects neurulation in mouse and is characterized by craniorachischisis, an open neural tube extending from the midbrain to the tail in embryos homozygous for the mutation. Lp maps to the distal portion of mouse chromosome 1, and as part of a positional cloning approach, we have generated a high-resolution linkage map of the Lp chromosomal region. For this, we have carried out extensive segregation analysis in a total of 706 backcross mice informative for Lp and derived from two crosses, (Lp/ + X SJL/J)F1 X SJL/J and (Lp/ + X SWR/J)F1 X SWR/J. In addition, 269 mice from a (Mus spretus X C57BL/6J)F1 X C57BL/6J interspecific backcross were also used to order marker loci and calculate intergene distances for this region. With these mice, a total of 28 DNA markers corresponding to either cloned genes or anonymous markers of the SSLP or SSCP-types were mapped within a 5-cM interval overlapping the Lp region, with the following locus order and interlocus distances (in cM): centromere-D1Mit110 / Atp1β1 / Cd3ζ / Cd3η / D1Mit145 — D1Hun14 / D1Mit15 — D1Mit111 / D1Mit112 — D1Mit114 — D1Mit148 / D1Mit205/ D1Mit36 / D1Mit146 / D1Mit147 / D1Mit270 / D1Hun13 — Fcgr2 — Mpp — Apoa2/Fcer1γ - Lp - D1Mit149 / Spna1/Fcer1α-Eph1-Hlx1/D1Mit62. These studies have allowed the delineation of a maximum genetic interval for Lp of 0.5 cM, a size amenable to physical mapping techniques.     

The organization of DNA sequences in the mouse genome

Analysis of the organization of nucleotide sequences in mouse genome is carried out on total DNA at different fragment size, reannealed to intermediate value of Cot, by Ag⁺-Cs₂SO₄ density gradient centrifugation. — According to nuclease S-1 resistance and kinetic renaturation curves mouse genome appears to be made up of non-repetitive DNA (76% of total DNA), middle repetitive DNA (average repetition frequency 2×10⁴ copies, 15% of total DNA), highly repetitive DNA (8% of total DNA) and fold-back DNA (renatured density 1.701 g/ml, 1% of total DNA).— Non-repetitive sequences are intercalated with short middle repetitive sequences. One third of non-repetitive sequences is longer than 4500 nucleotides, another third is long between 1800 and 4500 nucleotides, and the remainder is shorter than 1800 nucleotides. —Middle repetitive sequences are transcribed in vivo. The majority of the transcribed repeated sequences appears to be not linked to the bulk of non-repeated sequences at a DNA size of 1800 nucleotides. — The organization of mouse genome analyzed by Ag⁺-Cs₂SO₄ density gradient of reannealed DNA appears to be substantially different than that previously observed in human genome using the same technique.     

Behaviour and the concept of “Heritability” axioms of an ethological refutation

This paper discusses the widespread use of heritability calculations in recent behaviour research including behaviour genetics. In the sequel, a radical criticism concerning the basic axioms of the underlying, more general concept itself is presented. The starting point for testing the proclaimed universal validity of this concept stems from a fictitious yet realistic example taken from learning research. The theoretical result, based on the application of the conventional reasoning in this field, states that developmental processes — and learning is only one specific case out of an immense number of similar behavioural mechanisms — can neither be adequately described nor causally explained with sufficient reliability within the context of the heredity paradigm. On the contrary, an inherent inconsistency of the concept itself when applied to behaviour processes is demonstrated. Finally, a conceptual alternative involving a systems-theoretical approach to the problem is presented: In such a perspective it is the concept of cognition which represents the adequate explanatory theorem - a theorem in which quantitative processing of information from the environment is clearly revealed to belong to a subordinate level of living organization.     

How Random Is Social Behaviour? Disentangling Social Complexity through the Study of a Wild House Mouse Population

Out of all the complex phenomena displayed in the behaviour of animal groups, many are thought to be emergent properties of rather simple decisions at the individual level. Some of these phenomena may also be explained by random processes only. Here we investigate to what extent the interaction dynamics of a population of wild house mice (Mus domesticus) in their natural environment can be explained by a simple stochastic model. We first introduce the notion of perceptual landscape, a novel tool used here to describe the utilisation of space by the mouse colony based on the sampling of individuals in discrete locations. We then implement the behavioural assumptions of the perceptual landscape in a multi-agent simulation to verify their accuracy in the reproduction of observed social patterns. We find that many high-level features – with the exception of territoriality – of our behavioural dataset can be accounted for at the population level through the use of this simplified representation. Our findings underline the potential importance of random factors in the apparent complexity of the mice''s social structure. These results resonate in the general context of adaptive behaviour versus elementary environmental interactions.