COMMENTARIES. Ecology, evolution and division of labour in social insects
|
|
- Dinah Holland
- 8 years ago
- Views:
Transcription
1 Anim. Behav., 1997, 53, COMMENTARIES Ecology, evolution and division of labour in social insects JAMES F. A. TRANIELLO & REBECA B. ROSENGAUS Department of Biology, Boston University (Received 27 October 1994; initial acceptance 30 January 1995; final acceptance 18 March 1996; MS. number: AS-1149R) Caste and division of labour have formed the core of the study of the organization of insect societies for the past four decades. Indeed, the description and analysis of task allocation between colony members are fundamental to understanding the organization of a complex biological system whose functioning depends upon the behavioural integration of a potentially large number of individuals. Recent research has emphasized the dynamic nature and fluidity of task allocation (Gordon 1989, 1995), the role of self-organization (Page & Mitchell 1991) and the physiology and genetics of task partitioning (Page & Robinson 1991; Robinson 1992). The study of caste and division of labour, however, has historically been established in the evolution and ecology of social insects (Oster & Wilson 1978; Calabi & Traniello 1989; Hölldobler & Wilson 1990; Schmid-Hempel 1992; Tschinkel 1993; Beshers & Traniello 1994). Although the basic tenets of caste theory were first formulated to explain the origin and significance of physical castes, caste theory has also provided a conceptual framework to examine the adaptiveness of age-based division of labour, although this aspect of caste theory has rarely been tested and is thus poorly understood (Schmid-Hempel 1992). The theory predicts that the behavioural characteristics of individuals, including worker behavioural development, have been shaped as part of the overall evolutionary design of a colony of a given species. As Hölldobler & Wilson (1990, page 312) have stated, Each species has its own distinctive pattern of temporal polyethism. (In this commentary we consider the terms age-based division of labour and temporal polyethism to be interchangeable, and follow the definitions and Correspondence: J. F. A. Traniello, Department of Biology, Boston University, 5 Cummington Street, Boston, MA 02215, U.S.A. ( jft@bio.bu.edu). usage of Oster & Wilson (1978) and Hölldobler & Wilson (1990). We also believe that these terms and the social phenomena they describe adequately cover aspects of worker behavioural development and task performance flexibility, and therefore we prefer not to adopt Franks (1994) definition of temporal polyethism). The recent dialogue between Robinson et al. (1994) and Franks & Tofts (1994) provides a critical discussion of theoretical, genetic and physiological analyses of age-based division of labour; their debate on temporal polyethism relies on socially advanced hymenopteran species as model systems. The genetic and physiological approaches described by Robinson et al. (1994) focus on the mechanisms of polyethism and are relevant to the evolution and ecology of division of labour, because they detail the proximate basis of what appear to be adaptive patterns of behavioural development. Franks & Tofts (1994), in contrast, suggested that the behavioural schedules commonly identified as age polyethism emerge as a consequence of the rules workers follow in their search for tasks to perform in a colony. Specifically, they attempted to identify a simple regulatory process involving nest design and the spatial array of tasks to be carried out to explain polyethism (Tofts & Franks 1992; Sendova-Franks & Franks 1993; Franks & Tofts 1994; reviewed in Bourke & Franks 1995). Provocatively asserting that age polyethism is a myth and that the relationship between age and task performance is not causal, Franks (1994) advocated a paradigm shift in the study of polyethism. We argue, as have other researchers, that patterns of division of labour in the social insects appear to be highly variable and species-specific. And, as has long been recognized, we acknowledge that division of labour may be related to the /97/ $25.00/0/ar The Association for the Study of Animal Behaviour 209
2 210 Animal Behaviour, 53, 1 spatial array of tasks faced by a colony (Wilson 1976). The pattern of division of labour might depend upon worker development and reproductive potential, nest architecture, colony size, foraging ecology and other characteristics, and depending on the species, may or may not involve age-related behaviour and/or flexible aspects of task performance by workers (Oster & Wilson 1978; Wilson 1985). Interspecific variation in colony organization thus may be more accurately accounted for by historical and ecological factors than by a simple rule such as the foraging for work algorithm. Franks & Tofts (1994) and Tofts & Franks (1992) failed to acknowledge the diversity and significance of the individual and colony-level behavioural profiles that characterize different social insect species, and thus neglected the role of phylogeny as a constraint and the importance of ecology as a selective force in the organization of temporal division of labour. Given the diversity of social systems in insects, it would indeed be difficult for one organizational principle to have widespread phylogenetic application. In their rush to champion their model and herald what they claim as its broad significance, Tofts & Franks (1992) proceeded straight from the eusocial Hymenoptera to naked mole-rats without as much as a token evolutionary nod to the termites (Order Isoptera), a group phylogenetically unrelated to the Hymenoptera but strongly convergent in many aspects of sociality, including their systems of physical polymorphism and temporal polyethism. For example, the caste systems of the African fungus-growing termite Macrotermes parallel those of the Neotropical fungus-growing ant Atta (Wilson 1980; Bardertscher et al. 1983; Gerber et al. 1988). The socio-ecological similarity between ants and termites raises questions concerning the organization of division of labour and the applicability of the foraging for work to isopteran species. The foraging-for-work model assumes that the ordering of tasks by workers arises naturally from the structure of nests, and simplistically is just the distance at which the task can be performed from the centre of the brood pile (Tofts & Franks 1992, page 342; see also Tofts 1993). Termite species vary in nest structure and the degree of spatial segregation of tasks. Some termite species (the lower termites) have a onepiece nest, forming their colonies within their wood food source, but other species build elaborate mounds or carton structures and have a distinct food/nest separation (Abe 1987). One-piece-nest species, such as the dampwood termite, Zootermopsis angusticollis, provide an opportunity to examine division of labour in a species in which it is extremely easy to measure the spatial structure of some important colony activities. In this species, there appears to be no segregation of brood care and foraging, which in ants and higher termites occur inside and outside of the nest, respectively, because in Z. angusticollis the nest itself is the colony s food source. Yet there is a nest centre where the primary reproductives and eggs are found, and there appears to be a potential for task displacement because maturing larvae may eventually leave the labour force as they differentiate into alates and disperse. Developing larvae could move centrifugally from the nest centre to perform non-brood care tasks such as excavation, repair and feeding at the periphery of the nest and show some form of division of labour, but these termites seem to show no temporal polyethism (Rosengaus & Traniello 1993). First- and second-instar larvae are virtually inactive, and individuals from the third to seventh instars perform all colony labour with no apparent bias towards age or size. This study of termite polyethism was not designed as a specific test of the foraging for work algorithm, but results do offer a reasonable first approximation of the form of division of labour in a lower termite. The nutritional ecology of Z. angusticollis, among other factors, may provide insight into the social organization of this species (Rosengaus & Traniello 1993). Another concern in the evolution of polyethism in termites is that Z. angusticollis and other lower termites do not have a truly non-reproductive worker caste; larvae may eventually attain reproductive status in the parent nest (Noirot & Pasteels 1987). This reproductive plasticity could cause individuals to remain near the egg pile (where they might deposit their own eggs) and perform brood-related tasks as they age, rather than provide labour at other sites in the nest. West-Eberhard (1979, 1981) offered a similar argument concerning temporal polyethism in the social Hymenoptera, but suggested that in this group worker reproduction should result in a bias towards brood care in young adults that have functional ovaries. Because reproductive competency in termites increases with age, older larvae
3 Commentaries 211 may be predisposed to brood care in some isopteran species. Although insufficient information exists to permit conclusions about the effect of reproductive plasticity on task performance in the Isoptera, a comparison of the lower and higher termites suggests that worker sterility and temporal polyethism may be linked. As discussed above, termites in the lower sub-families construct onepiece nests; an individual termite retains the ability to reproduce through its life. In at least one of these species, labour does not appear to be organized through temporal castes (Rosengaus & Traniello 1993). Termite species in the higher subfamilies have separate nesting and feeding habits, a true sterile worker caste and welldeveloped age-related division of labour (Jones 1980; Abe 1987; Noirot & Pasteels 1987; Gerber et al. 1988; Higashi et al. 1991). The loss of reproductive options among workers and/or the separation of nest from food source therefore appear to have been prerequisites for the evolution of termite polyethism. Perhaps the separation of nest and food source created a spatial predisposition for the centrifugal movement of workers through task space as predicted by Tofts & Franks model. However, age-related behaviour in termites may have depended on nest stability and the evolution of worker sterility, rather than emerging as an inevitable by-product of workers foraging for work. A comprehensive explanatory framework for the evolution of temporal division of labour in termites thus appears to require an understanding of termite reproductive biology and nesting ecology. In addition, the so-called primitive ant species appear to have biological characteristics that offer important insights into their patterns of labour organization. Moreover, some species have attributes that appear to satisfy the assumptions of the foraging-for-work model but do not seem to provide support for it. The ponerine ant Amblyopone pallipes is a case in point. This species lacks age-based division of labour (Traniello 1978), yet has a nest structure that is sufficiently complex to generate the pattern of polyethism predicted by Tofts & Franks algorithm. In contrast to most ants, workers begin to forage after eclosion. In A. pallipes, foraging is part of brood-care behaviour because larvae are directly provisioned with prey, and the same worker performs both tasks. The Tofts & Franks task displacement model does not explain this atypical pattern of task performance in A. pallipes and other primitive ants as effectively as arguments that concern demographic constraints, foraging specialization and the retention of ancestral behavioural traits (Traniello 1978; Traniello & Jayasuriya 1985; Jaisson et al. 1992). Also, the degree of sociality itself may play a role in generating division of labour in A. pallipes and other ancestral ant species. For example, Nothomyrmecia macrops is considered to be the closest among living ant species to the ancestral ants; individuals infrequently interact in N. macrops colonies, and no age-based division of labour exists, although colonies have a nest centre with brood, and workers forage outside the nest (Jaisson et al. 1992). At what level of social complexity should we expect tasks to allocate workers? We do not draw attention to the biology of the above-named species simply to illustrate curious variation and an apparent lack of congruence with the foraging-for-work model, but to accentuate the need for a comparative socioecological approach in the study of division of labour. Robinson et al. (1994) noted that behaviour is best understood by integrating analyses at multiple levels of biological organization. In the case of division of labour, the results obtained at different levels should be synthesized and reconciled in light of the evolution and ecology of each species to form an understanding of the how and why of polyethism. There is strong evidence in favour of a causal relationship between worker age and task performance (Calderone 1995) and there is ample documentation of the genetic and hormonal correlates of behavioural development in some social insect species (Page & Robinson 1991; Robinson 1992; Robinson et al. 1994). There is also evidence to show that behavioural development is flexible and task allocation is socially regulated (Hölldobler & Wilson 1990; Gordon 1995). Empirical research on the ecology of division of labour indicates that worker age distributions are not necessarily associated with task schedules, and that the demographic and flexible behavioural components that contribute to colony-level patterns of division of labour may be selected over evolutionary and ecological time scales (Calabi & Traniello 1989). Realistic models and simulations and developmental studies of task allocation can
4 212 Animal Behaviour, 53, 1 complement physiological and etho-ecological approaches and perhaps offer alternative hypotheses to explain temporal polyethism. To fully achieve the integrative approach that is of current interest in the study of biological complexity, however, the role of ecology and historical constraints must be recognized. We believe that such a biological approach will continue to lead the search for general principles of the organization of division of labour. Describing the autonomy of biology in relation to the physical sciences, Mayr (1982, page 36) suggested that biologists should view the application of physical models to biological systems with great caution. He noted that when in the past naturalists and other biologists have stressed the importance of uniqueness and history in biology, their efforts were erroneously brushed aside as bad science by physical scientists and that the biological approach was considered inferior because it lacked mathematical form. We do not doubt that simple models can be valuable in understanding the dynamics of a complex system such as an insect society, if one is aware that the system itself is biological and is likely to be more complex and varied than a model. We thank the Whitehall Foundation and the National Science Foundation for research support, Sam Beshers, Robert L. Jeanne, Gene Robinson, Simon Robson and Barbara Thorne for discussion, and Fred Dyer and two anonymous referees for their thoughtful comments. REFERENCES Abe, T Evolution of life types in termites. In: Evolution and Coadaptation in Biotic Communities (Ed. by S. Kawano, J. H. Connell & T. Hidaka), pp Tokyo: University of Tokyo Press. Bardertscher, S., Gerber, C. & Leuthold, R. H Polyethism in food supply and processing in termite colonies of Macrotermes subhyalinus (Isoptera). Behav. Ecol. Sociobiol., 12, Beshers, S. & Traniello, J. F. A The adaptiveness of demographic distributions of the fungusgrowing ant Trachymyrmex septentrionalis. Ecology, 75, Bourke, A. F. G. & Franks, N. R Social Evolution in Ants. Princeton, New Jersey: Princeton University Press. Calabi, P. & Traniello, J. F. A Social organization in the ant Pheidole dentata: physical and age caste ratios lack ecological correlates. Behav. Ecol. Sociobiol., 24, Calderone, N. W Temporal division of labor in the honey bee, Apis mellifera: a developmental process or the result of environmental influences? Can. J. Zool., 73, Franks, N. R. & Tofts, C Foraging for work: how tasks allocate workers. Anim. Behav., 48, Gerber, C., Bardertscher, S. & Leuthold, R. H Polyethism in Macrotermes bellicosus (Isoptera). Insectes soc., 35, Gordon, D. M Dynamics of task switching in harvester ants. Anim. Behav., 38, Gordon, D. M The development of organization in an ant colony. Am. Scient., 83, Higashi, M., Yamamura, N., Abe, T. & Burns, T. P Why don t all termite species have a sterile worker caste? Proc. R. Soc. Ser. B, 246, Hölldobler, B. & Wilson, E. O The Ants. Cambridge, Massachusetts: Harvard University Press. Jaisson, P., Fresneau, D., Taylor, R. W. & Lenoir, A Social organization in some primitive ants. I. Nothomyrmecia macrops Clark. Insectes soc., 39, Jones, R. J Gallery construction by Nasutitermes costalis: polyethism and the behavior of individuals. Insectes soc., 27, Mayr, E The Growth of Biological Thought. Cambridge, Massachusetts: Harvard University Press. Noirot, C. & Pasteels, J. M Ontogenic development and evolution of the worker caste in termites. Experientia, 43, Oster, G. F. & Wilson, E. O Caste and Ecology in the Social Insects. Princeton, New Jersey: Princeton University Press. Page, R. E., Jr & Mitchell, S. D Self organization and adaptation in insect societies. In: PSA 1990, Vol. 2 (Ed. by A. Fine, M. Forbes & L. Wessels), pp East Lansing, Michigan: Philosophical Science Association. Page, R. E., Jr & Robinson, G. E The genetics of division of labour in honeybee colonies. Adv. Insect Physiol., 35, Robinson, G. E Regulation of division of labor in insect societies. A. Rev. Entomol., 37, Robinson, G. E., Page, R. E., Jr & Huang, Z.-Y Temporal polyethism in social insects is a developmental process. Anim. Behav., 48, Rosengaus, R. B. & Traniello, J. F. A Temporal polyethism in incipient colonies of the dampwood termite Zootermopsis angusticollis: a single multiage caste. J. Insect Behav., 6, Schmid-Hempel, P Worker castes and adaptive demography. J. evol. Biol., 5, Sendova-Franks, A. & Franks, N. R Task allocation in ant colonies within variable environments (a study of temporal polyethism: experimental). Bull. math. Biol., 55, Tofts, C Algorithms for task allocation in ants (a study of temporal polyethism: theory). Bull. math. Biol., 55, Tofts, C. & Franks, N. R Doing the right thing: ants, honeybees and naked mole rats. Trends Ecol. Evol., 7,
5 Commentaries 213 Traniello, J. F. A Caste in a primitive ant: absence of age polyethism in Amblyopone. Science, 202, Traniello, J. F. A. & Jayasuriya, A. K The biology of the primitive ant Aneuretus simoni (Emery). II. The social ethogram and division of labor. Insectes soc., 32, Tschinkel, W. R Sociometry and sociogenesis of colonies of the fire ant Solenopsis invicta during one annual cycle. Ecol. Monogr., 63, West-Eberhard, M. J Sexual selection, social competition, and evolution. Proc. Am. Phil. Soc., 123, West-Eberhard, M. J Intragroup selection and the evolution of insect societies. In: Natural Selection and Social Behavior (Ed. by R. D. Alexander & D. W. Tinkle), pp New York: Chiron Press. Wilson, E. O Behavioral discretization and the number of castes in an ant species. Behav. Ecol. Sociobiol., 1, Wilson, E. O Caste and division of labor in leaf-cutter ants (Hymenoptera: Formicidae: Atta). I. The overall pattern in A. sexdens. Behav. Ecol. Sociobiol., 7, Wilson, E. O The sociogenesis of insect colonies. Science, 228,
Castes in Social Insects
Castes in Social Insects Module Chemical Ecology, Ayasse Outline Castes in Social Insects 1. definition 2. castes in social insects 3. caste determination A. psychophysiological caste determination B.
More informationSocial Insects. Social Insects. Subsocial 4/11/10. More widespread 13 orders of insects no reproductive division of labor
Social Insects Sociality evolved multiple times in insects Much of Earth s fauna consists of social insects They play major roles in entire ecosystems Proliferation of ants and termites associated with
More informationBehavioral Repertory of the Weaver Ant Camponotus (Myrmobrachys) senex (Hymenoptera: Formicidae)
Behavioral Repertory of the Weaver Ant Camponotus (Myrmobrachys) senex (Hymenoptera: Formicidae) by Jean Carlos Santos, Marcela Yamamoto, Flávio Rodrigues Oliveira & Kleber Del-Claro 1 ABSTRACT Weaver
More informationOn foraging, recruitment systems and optimum number of scouts in eusocial colonies
Ins. Soc. 39:201-213 (1992) 1015-1621/92/020201-13 $1.50 + 0.20/0 9 1992 Birkh/iuser Verlag, Basel On foraging, recruitment systems and optimum number of scouts in eusocial colonies K. Jaffe 1 and J. L.
More informationIntroduction. Swarm Intelligence - Thiemo Krink EVALife Group, Dept. of Computer Science, University of Aarhus
Swarm Intelligence - Thiemo Krink EVALife Group, Dept. of Computer Science, University of Aarhus Why do we need new computing techniques? The computer revolution changed human societies: communication
More informationSTUDENT VERSION INSECT COLONY SURVIVAL OPTIMIZATION
STUDENT VERSION INSECT COLONY SURVIVAL OPTIMIZATION STATEMENT We model insect colony propagation or survival from nature using differential equations. We ask you to analyze and report on what is going
More informationSymbiont-derived -1,3-glucanases in a social insect: mutualism beyond nutrition
Electronic Supplementary Material Frontiers in Microbiology Microbial Symbioses Original Research Article Symbiont-derived -1,3-glucanases in a social insect: mutualism beyond nutrition Rebeca B. Rosengaus
More informationNext Generation Science Standards
The Next Generation Science Standards and the Life Sciences The important features of life science standards for elementary, middle, and high school levels Rodger W. Bybee Publication of the Next Generation
More informationOkami Study Guide: Chapter 3 1
Okami Study Guide: Chapter 3 1 Chapter in Review 1. Heredity is the tendency of offspring to resemble their parents in various ways. Genes are units of heredity. They are functional strands of DNA grouped
More informationAP Biology Essential Knowledge Student Diagnostic
AP Biology Essential Knowledge Student Diagnostic Background The Essential Knowledge statements provided in the AP Biology Curriculum Framework are scientific claims describing phenomenon occurring in
More informationCharles Elton once remarked that All cold-blooded animals
Behavioral Ecology Vol. 12 No. 1: 111 119 The adaptive value of inactive foragers and the scout-recruit system in honey bee (Apis mellifera) colonies Carl Anderson Department of Genetics and Ecology, University
More informationAnimal Models of Human Behavioral and Social Processes: What is a Good Animal Model? Dario Maestripieri
Animal Models of Human Behavioral and Social Processes: What is a Good Animal Model? Dario Maestripieri Criteria for assessing the validity of animal models of human behavioral research Face validity:
More informationPS3021, PS3022, PS4040
School of Psychology Important Degree Information: B.Sc./M.A. Honours The general requirements are 480 credits over a period of normally 4 years (and not more than 5 years) or part-time equivalent; the
More informationDriver Ants Invading a Termite Nest: Why Do the Most Catholic Predators of All Seldom Take This Abundant Prey?
BIOTROPICA 39(5): 663 667 2007 10.1111/j.1744-7429.2007.00296.x Driver Ants Invading a Termite Nest: Why Do the Most Catholic Predators of All Seldom Take This Abundant Prey? Caspar Schöning 1 Institute
More informationBiological kinds and the causal theory of reference
Biological kinds and the causal theory of reference Ingo Brigandt Department of History and Philosophy of Science 1017 Cathedral of Learning University of Pittsburgh Pittsburgh, PA 15260 E-mail: inb1@pitt.edu
More informationG u i d e l i n e s f o r K12 Global C l i m a t e Change Education
G u i d e l i n e s f o r K12 Global C l i m a t e Change Education Adapted by: by the National Wildlife Federation from the Environmental Education Guidelines for Excellence of the North American Association
More informationTermite Behavior: Focus on Experimental Design and Results (adapted from Surmacz, 2004 and Matthews and Matthews, 2012)
Termite Behavior: Focus on Experimental Design and Results (adapted from Surmacz, 2004 and Matthews and Matthews, 2012) Prior to coming to lab this week, you should read all of the introductory information.
More informationDr V. J. Brown. Neuroscience (see Biomedical Sciences) History, Philosophy, Social Anthropology, Theological Studies.
Psychology - pathways & 1000 Level modules School of Psychology Head of School Degree Programmes Single Honours Degree: Joint Honours Degrees: Dr V. J. Brown Psychology Neuroscience (see Biomedical Sciences)
More informationUnifying Epistemologies by Combining World, Description and Observer
Unifying Epistemologies by Combining World, Description and Observer Stuart Umpleby Research Program in Social and Organizational Learning The George Washington University Washington, DC Umpleby@gwu.edu
More informationPSYCHOLOGY PROGRAM LEARNING GOALS, LEARNING OUTCOMES AND COURSE ALLIGNMENT MATRIX. 8 Oct. 2010
PSYCHOLOGY PROGRAM LEARNING GOALS, LEARNING OUTCOMES AND COURSE ALLIGNMENT MATRIX 8 Oct. 2010 Departmental Learning Goals and Outcomes LEARNING GOAL 1: KNOWLEDGE BASE OF PSYCHOLOGY Demonstrate familiarity
More informationTECHNICALPUBLICATION TERMITES IN NEW SOUTH WALES PART 1. TERMITE BIOLOGY. MA Horwood and RH Eldridge TERMITE COLONIES
TECHNICALPUBLICATION FOREST RESOURCES RESEARCH NUMBER 21 ISSN: 0155-7548 TERMITES IN NE SOUTH ALES PART 1. TERMITE BIOLOGY MA Horwood and RH Eldridge Termites are pallid, secretive creatures colloquially
More informationTask Allocation in Ant Colonies
Task Allocation in Ant Colonies Alejandro Cornejo 1, Anna Dornhaus 2, Nancy Lynch 3, and Radhika Nagpal 1 1 Harvard University School of Engineering and Applied Sciences 2 University of Arizona Ecology
More informationBridging Micro and Macro Domains: Workforce Differentiation and Strategic Human Resource Management
Special Issue: Bridging Micro and Macro Domains Journal of Management Vol. 37 No. 2, March 2011 421-428 DOI: 10.1177/0149206310373400 The Author(s) 2011 Reprints and permission: http://www. sagepub.com/journalspermissions.nav
More informationCollege of Agriculture
100 INTRODUCTION TO ESTRY. (3) A brief coverage of the general fields of forestry; development and importance; tree growth; principal forest regions and important timber species; forest management practices;
More informationPSYCHOLOGY PROGRAM LEARNING GOALS AND OUTCOMES BY COURSE LISTING
PSYCHOLOGY PROGRAM LEARNING GOALS AND OUTCOMES BY COURSE LISTING Psychology 1010: General Psychology Learning Goals and Outcomes LEARNING GOAL 1: KNOWLEDGE BASE OF PSYCHOLOGY Demonstrate familiarity with
More informationUndergraduate Psychology Major Learning Goals and Outcomes i
Undergraduate Psychology Major Learning Goals and Outcomes i Goal 1: Knowledge Base of Psychology Demonstrate familiarity with the major concepts, theoretical perspectives, empirical findings, and historical
More informationCollege of Arts and Sciences: Social Science and Humanities Outcomes
College of Arts and Sciences: Social Science and Humanities Outcomes Communication Information Mgt/ Quantitative Skills Valuing/Ethics/ Integrity Critical Thinking Content Knowledge Application/ Internship
More informationBug Power. Amazing Ants. Busy Bees. Teamwork. How do some insects work together?
Non-fiction: Bug Power Bug Power Teamwork How do some insects work together? What do termites, ants, and honeybees have in common? They are all social (SOHshuhl) insects. Social insects live together in
More informationHidden Aggression in Termite Workers: Plastic. Defensive Behaviour Dependent upon Social Context
Title Hidden aggression in termite worker behaviour dependent upon social con Author(s) Ishikawa, Yuki; Miura, Toru Citation Animal Behaviour, 83(3): 737-745 Issue Date 2012-03 DOI Doc URLhttp://hdl.handle.net/2115/48599
More informationCPO Science and the NGSS
CPO Science and the NGSS It is no coincidence that the performance expectations in the Next Generation Science Standards (NGSS) are all action-based. The NGSS champion the idea that science content cannot
More informationNegative Effects of Azteca Ants on the Distribution of the Termite Neocapritermes braziliensis in Central Amazonia ABSTRACT INTRODUCTION
Negative Effects of Azteca Ants on the Distribution of the Termite Neocapritermes braziliensis in Central Amazonia by Pedro A. C. Lima Pequeno 1* & Pauline O. Pantoja 1 ABSTRACT Termites play important
More informationSoft Skills Requirements in Software Architecture s Job: An Exploratory Study
Soft Skills Requirements in Software Architecture s Job: An Exploratory Study 1 Faheem Ahmed, 1 Piers Campbell, 1 Azam Beg, 2 Luiz Fernando Capretz 1 Faculty of Information Technology, United Arab Emirates
More informationUtah State Office of Education Elementary STEM Endorsement Course Framework Nature of Science and Engineering
Course Description: Utah State Office of Education Elementary STEM Endorsement Course Framework Nature of Science and Engineering In this course participants will experience introductory explorations of
More informationProgram in Ecology, Evolution and Conservation Biology, Urbana, IL Ph.D. in progress Advisor: Gene E. Robinson
Beryl M. Jones Program in Ecology, Evolution, and Conservation Biology University of Illinois at Urbana-Champaign 286 Morrill Hall, 505 South Goodwin Avenue Urbana, IL 61801 EDUCATION University of Illinois
More informationWeb Mining using Artificial Ant Colonies : A Survey
Web Mining using Artificial Ant Colonies : A Survey Richa Gupta Department of Computer Science University of Delhi ABSTRACT : Web mining has been very crucial to any organization as it provides useful
More informationCurriculum Vitae. Teresa L. McElhinny
Curriculum Vitae Teresa L. McElhinny 4588 Van Atta Rd Okemos, MI 48864 517 420 4288 mcelhinn@msu.edu http://terrimac.com/ Education PhD 2009, Departments of Zoology and Ecology, Evolutionary Biology &
More informationNOTE: This document is specific to the 2000-2002 printed catalog.
NOTE: This document is specific to the 2000-2002 printed catalog. H BACHELOR OF SCIENCE MINOR PROGRAM DESCRIPTION The academic area of study in Health Science is part of the total program of the College
More informationBOSTON UNIVERSITY SCHOOL OF PUBLIC HEALTH PUBLIC HEALTH COMPETENCIES
BOSTON UNIVERSITY SCHOOL OF PUBLIC HEALTH PUBLIC HEALTH COMPETENCIES Competency-based education focuses on what students need to know and be able to do in varying and complex situations. These competencies
More informationKing, B.H. and S.W. Skinner. 1991. Sex ratio in a new species of Nasonia with fully-winged males. Evolution 45:225-228.
King, B.H. and S.W. Skinner. 1991. Sex ratio in a new species of Nasonia with fully-winged males. Evolution 45:225-228. This version matches the text but not the formatting of the published article. SEX
More informationPsychology. 42 Credits Complete the requirements shown in the General Education Requirements section of this catalog. Include this specific course.
Psychology Psychology is the science of behavior and mental processes. Behavior is anything an organism does that we can observe and record; examples include smiling, talking, yelling, and marking a questionnaire.
More informationCenter for Effective Organizations
Center for Effective Organizations WHAT MAKES HR A STRATEGIC PARTNER? CEO PUBLICATION G 09-01 (555) EDWARD E. LAWLER III Center for Effective Organizations Marshall School of Business University of Southern
More informationIntroduction to 30th Anniversary Perspectives on Cognitive Science: Past, Present, and Future
Topics in Cognitive Science 2 (2010) 322 327 Copyright Ó 2010 Cognitive Science Society, Inc. All rights reserved. ISSN: 1756-8757 print / 1756-8765 online DOI: 10.1111/j.1756-8765.2010.01104.x Introduction
More informationPsychology (MA) Program Requirements 36 credits are required for the Master's Degree in Psychology as follows:
Psychology (MA) ACADEMIC DIRECTOR: Carla Marquez-Lewis CUNY School of Professional Studies 101 West 31 st Street, 7 th Floor New York, NY 10001 Email Contact: Carla Marquez-Lewis, carla.marquez-lewis@cuny.edu
More informationHistory of the study of animal behavior
History of the study of animal behavior 100,000 years B.P. through ~1850 People have been making detailed descriptions of the natural history of animals since the dawn of our species 100,000 years B.P.
More informationThe primary goal of this thesis was to understand how the spatial dependence of
5 General discussion 5.1 Introduction The primary goal of this thesis was to understand how the spatial dependence of consumer attitudes can be modeled, what additional benefits the recovering of spatial
More informationIt is widely accepted by those in the scientific community that women have been
1 It is widely accepted by those in the scientific community that women have been systematically disregarded and discriminated against in science. However, the extent to which this has undermined aspirations
More informationCHAOS, COMPLEXITY, AND FLOCKING BEHAVIOR: METAPHORS FOR LEARNING
GIVING WINGS TO NEW IDEAS Wingspread Journal, Summer 1996 CHAOS, COMPLEXITY, AND FLOCKING BEHAVIOR: METAPHORS FOR LEARNING by Stephanie Pace Marshall Sir Isaac Newton saw the universe as an orderly clock.
More informationPractice Questions 1: Evolution
Practice Questions 1: Evolution 1. Which concept is best illustrated in the flowchart below? A. natural selection B. genetic manipulation C. dynamic equilibrium D. material cycles 2. The diagram below
More informationKnowledge: The successful individual in the sport management profession will:
268 SPORT MANAGEMENT Sport Management Program Outcomes Knowledge: The successful individual in the sport management profession will: 1. Understand and apply the principles of health, physical fitness and
More informationSchool of Advanced Studies Doctor Of Education In Educational Leadership With A Specialization In Educational Technology. EDD/ET 003 Requirements
School of Advanced Studies Doctor Of Education In Educational Leadership With A Specialization In Educational Technology The mission of the Doctor of Education in Educational Leadership degree program
More informationCourse Name Social Psychology
NPTEL Online - IIT Kanpur Course Name Social Psychology Department Department of Humanities and Social Sciences IIT Kanpur Instructor Dr. Kumar Ravi Priya file:///c /Documents%20and%20Settings/iitkrana1/Desktop/Social%20Psychology/lecture1/main.html[5/1/2014
More informationPsychology. Administered by the Department of Psychology within the College of Arts and Sciences.
Psychology Dr. Spencer Thompson, Professor, is the Chair of Psychology and Coordinator of Child and Family Studies. After receiving his Ph.D. in Developmental Psychology at the University of California,
More informationProgram Approval Form
Program Approval Form For approval of new programs and deletions or modifications to an existing program. Action Requested: Type (Check one): Create New (SCHEV approval required except for minors) B.A.
More informationStable matching: Theory, evidence, and practical design
THE PRIZE IN ECONOMIC SCIENCES 2012 INFORMATION FOR THE PUBLIC Stable matching: Theory, evidence, and practical design This year s Prize to Lloyd Shapley and Alvin Roth extends from abstract theory developed
More informationI. Insect sociality II. Evolution of Eusociality (ants and termites) III. Success of Eusocial Insects IV. Ants V. Ant plants.
I. Insect sociality II. Evolution of Eusociality (ants and termites) III. Success of Eusocial Insects IV. Ants V. Ant plants Social Insects Degrees of Sociality Solitary: exhibit no social behavior - most
More informationA Survey on Load Balancing Techniques Using ACO Algorithm
A Survey on Load Balancing Techniques Using ACO Algorithm Preeti Kushwah Department of Computer Science & Engineering, Acropolis Institute of Technology and Research Indore bypass road Mangliya square
More informationWhat is Reflective Practice? Joy Amulya Senior Associate Community Science
What is Reflective Practice? Joy Amulya Senior Associate Community Science Community Science 0 May 2011 What is Reflective Practice? Joy Amulya Senior Associate, Community Science Reflection: the foundation
More informationWhat mathematical optimization can, and cannot, do for biologists. Steven Kelk Department of Knowledge Engineering (DKE) Maastricht University, NL
What mathematical optimization can, and cannot, do for biologists Steven Kelk Department of Knowledge Engineering (DKE) Maastricht University, NL Introduction There is no shortage of literature about the
More informationProfessor S D Reicher. Needs. Needs. Adults with Learning Disabilities who have Significant and Complex Needs. Health Psychology
School of Psychology Head of School Degree Programmes Postgraduate Certificate: Professor S D Reicher Adults with Learning Disabilities who have Significant and Complex Needs Postgraduate Diploma: Adults
More informationContent Delivery Network (CDN) and P2P Model
A multi-agent algorithm to improve content management in CDN networks Agostino Forestiero, forestiero@icar.cnr.it Carlo Mastroianni, mastroianni@icar.cnr.it ICAR-CNR Institute for High Performance Computing
More informationTHE REASONING ART: or, The Need for an Analytical Theory of Architecture
P ROCEEDINGS VOLUME I SPACE SYNTAX TODAY THE REASONING ART: or, The Need for an Analytical Theory of Architecture Professor Bill Hillier and Dr Julienne Hanson University College London, London, England
More informationA Service Revenue-oriented Task Scheduling Model of Cloud Computing
Journal of Information & Computational Science 10:10 (2013) 3153 3161 July 1, 2013 Available at http://www.joics.com A Service Revenue-oriented Task Scheduling Model of Cloud Computing Jianguang Deng a,b,,
More informationVygotsky s Theory: Lessons for Student Engagement Research
SERU International Research Conference, May15-17, 2014 Vygotsky s Theory: Lessons for Student Engagement Research Natalia Maloshonok National Research University Higher School of Economics E-mail: nmaloshonok@hse.ru
More informationEvolutionary Psychology as a Metatheory for the Social Sciences
Evolutionary Psychology as a Metatheory for the Social Sciences Annemie Ploeger 1 Abstract: Evolutionary psychology has been proposed as a metatheory for the social sciences. In this paper, the different
More informationCritical thinking (7 courses total, 5 Foundations and 2 Applications) An illustration of how the categories of area of the grid could be developed
Critical thinking (7 courses total, 5 Foundations and 2 Applications) An illustration of how the categories of area of the grid could be developed Critical thinking is the starting point and context of
More informationGRADUATE CATALOG LISTING
GRADUATE CATALOG LISTING 1 BIOINFORMATICS & COMPUTATIONAL BIOLOGY Telephone: (302) 831-0161 http://bioinformatics.udel.edu/education Faculty Listing: http://bioinformatics.udel.edu/education/faculty A.
More informationECOLOGICAL RESTORATION A MEANS OF CONSERVING BIODIVERSITY AND SUSTAINING LIVELIHOODS
ECOLOGICAL A MEANS OF CONSERVING BIODIVERSITY AND SUSTAINING LIVELIHOODS RESTORATION The Society for Ecological Restoration International (SER) is a non-profit organization infused with the energy of involved
More informationMARK SCHEME for the May/June 2007 question paper 9699 SOCIOLOGY. 9699/02 Paper 2 (Data Response), maximum raw mark 50
www.xtremepapers.com UNIVERSITY OF CAMBRIDGE INTERNATIONAL EXAMINATIONS GCE Advanced Subsidiary Level and GCE Advanced Level MARK SCHEME for the May/June 2007 question paper 9699 SOCIOLOGY 9699/02 Paper
More informationBehavioral Sciences INDIVIDUAL PROGRAM INFORMATION 2015 2016. 866.Macomb1 (866.622.6621) www.macomb.edu
Behavioral Sciences INDIVIDUAL PROGRAM INFORMATION 2015 2016 866.Macomb1 (866.622.6621) www.macomb.edu Behavioral Sciences CREDENTIAL TITLE PROGRAM OPTIONS CREDIT HOURS REQUIRED Certificate Behavioral
More informationProblems With Programmable self-assembly in a thousand-robot swarm
Problems With Programmable self-assembly in a thousand-robot swarm Affiliations: Department of Computer Science, COMSATS Institute of IT, Islamabad, Pakistan Author: Muaz A. Niazi* ǂ *Correspondence to:
More informationA CONTENT STANDARD IS NOT MET UNLESS APPLICABLE CHARACTERISTICS OF SCIENCE ARE ALSO ADDRESSED AT THE SAME TIME.
Environmental Science Curriculum The Georgia Performance Standards are designed to provide students with the knowledge and skills for proficiency in science. The Project 2061 s Benchmarks for Science Literacy
More informationBiology of Termites: A Modern Synthesis
Biology of Termites: A Modern Synthesis David Edward Bignell Yves Roisin Nathan Lo Editors Biology of Termites: A Modern Synthesis Foreword by Bernard J. Crespi 123 Editors Prof. David Edward Bignell
More informationYour Questions from Chapter 1. General Psychology PSYC 200. Your Questions from Chapter 1. Your Questions from Chapter 1. Science is a Method.
General Psychology PSYC 200 Methods of Psychology Your Questions from Chapter 1 Which names of people are important to remember? In what way are we going to be tested on this material? What exactly did
More informationBIOLOGICAL SCIENCES REQUIREMENTS [63 75 UNITS]
Biological Sciences Major The Biological Sciences address many of the most important and fundamental questions about our world: What is life? How does our brain produce our ideas and emotions? What are
More informationby Chow-Yang Lee 1, Charunee Vongkaluang 2 & Michael Lenz 3
213 Challenges to Subterranean Termite Management of Multi-Genera Faunas In Southeast Asia and Australia by Chow-Yang Lee 1, Charunee Vongkaluang 2 & Michael Lenz 3 Abstract Termites are an important group
More informationMETAFLUMIZONE: A NEW INSECTICIDE FOR URBAN INSECT CONTROL FROM BASF
Proceedings of the Fifth International Conference on Urban Pests Chow-Yang Lee and William H. Robinson (editors), 2005. Printed by Perniagaan Ph ng @ P&Y Design Network, Malaysia. METAFLUMIZONE: A NEW
More informationThe Role of Computers in Synchronous Collaborative Design
The Role of Computers in Synchronous Collaborative Design Wassim M. Jabi, The University of Michigan Theodore W. Hall, Chinese University of Hong Kong Abstract In this paper we discuss the role of computers
More information2016 Bee College Tentative Short Course Descriptions
2016 Bee College Tentative Short Course Descriptions A Beekeeper s Year For starting beekeepers, the first year could be the hardest since you are trying to figure out all the quirks of beekeeping. Come
More informationAndrea Gilli, PhD Post-doctoral Fellow - Center for Security Studies Metropolitan University Prague
Andrea Gilli, PhD Post-doctoral Fellow - Center for Security Studies Metropolitan University Prague PH.D. DISSERTATION EXECUTIVE SUMMARY Unipolarity, Technological Change and Arms Manufacturing: Industrial
More informationNEUROEVOLUTION OF AUTO-TEACHING ARCHITECTURES
NEUROEVOLUTION OF AUTO-TEACHING ARCHITECTURES EDWARD ROBINSON & JOHN A. BULLINARIA School of Computer Science, University of Birmingham Edgbaston, Birmingham, B15 2TT, UK e.robinson@cs.bham.ac.uk This
More informationBRAESS-LIKE PARADOXES FOR NON-COOPERATIVE DYNAMIC LOAD BALANCING IN DISTRIBUTED COMPUTER SYSTEMS
GESJ: Computer Science and Telecommunications 21 No.3(26) BRAESS-LIKE PARADOXES FOR NON-COOPERATIVE DYNAMIC LOAD BALANCING IN DISTRIBUTED COMPUTER SYSTEMS Said Fathy El-Zoghdy Department of Computer Science,
More informationUnderstanding by Design. Title: BIOLOGY/LAB. Established Goal(s) / Content Standard(s): Essential Question(s) Understanding(s):
Understanding by Design Title: BIOLOGY/LAB Standard: EVOLUTION and BIODIVERSITY Grade(s):9/10/11/12 Established Goal(s) / Content Standard(s): 5. Evolution and Biodiversity Central Concepts: Evolution
More informationEvaluating the Lead Time Demand Distribution for (r, Q) Policies Under Intermittent Demand
Proceedings of the 2009 Industrial Engineering Research Conference Evaluating the Lead Time Demand Distribution for (r, Q) Policies Under Intermittent Demand Yasin Unlu, Manuel D. Rossetti Department of
More informationA Preliminary Proposal for a Pharmaceutical Engineering Graduate Program
A Preliminary Proposal for a Pharmaceutical Engineering Graduate Program Planning Committee: Prabir Basu Steve Byrn Ken Morris Rex Reklaitis Paul Sojka Venkat Venkatasubramanian Carl Wassgren National
More information1998 B.S. Bioenvironmental Science and Entomology, Texas A&M University, College Station, Texas.
Janis J. Reed Extension Program Specialist Texas A&M AgriLife Extension 2488 TAMU College Station, TX 77843-2488 Email: janisreed@tamu.edu Phone: 512-970-3904 Education 2013 Ph.D. Entomology Texas A&M
More informationAnimal behavior can be investigated in a variety of
7 Behavioral Ecology Concept Outline 7. Evolutionary forces shape behavior. Behavioral Ecology. Behavior is shaped by natural selection. Foraging Behavior. Natural selection favors the most efficient foraging
More informationEcological models in chemical risk assessment Preliminary Recommendations of the SETAC Europe workshop MODELINK
Ecological models in chemical risk assessment Preliminary Recommendations of the SETAC Europe workshop MODELINK V. Ducrot, A. Alix, D. Auteri, P. Carpentier, P. Dohmen, V. Forbes, V. Grimm, U. Hommen,T.G.
More informationPSYC PSYCHOLOGY. 2011-2012 Calendar Proof
PSYC PSYCHOLOGY PSYC1003 is a prerequisite for PSYC1004 and PSYC1004 is a prerequisite for all remaining Psychology courses. Note: See beginning of Section F for abbreviations, course numbers and coding.
More informationMS. Structure, Function, and Information Processing
MIDDLE SCHOOL LIFE SCIENCE Alignment with National Science Standards Use the chart below to find Science A-Z units that best support the Next Generation Science Standards* for Middle School Life Science,
More informationCurrently 32 million, Canada s population is
Geoff Bowlby This paper was originally presented to the Paris Group, a body of statisticians and economists from national statistical organizations around the world interested in improving labour statistics.
More informationMaster s Entry into Nursing. Academic Manual 2015-2016
Master s Entry into Nursing Academic Manual 2015-2016 TABLE OF CONTENTS Overview of the Master s Entry into Nursing (MEN) Program 2 Outcomes 2 Plan of Study 3-4 Course Descriptions 5-11 Overview of the
More informationUSING STARLOGO TO INTRODUCE DIFFERENTIAL EQUATIONS
USING STARLOGO TO INTRODUCE DIFFERENTIAL EQUATIONS PHILIP ANDERSON AND CARL R. SEAQUIST Abstract. Massively parallel programming languages, like StarLogo, provide a rich environment for introducing differential
More informationDepartment: PSYC. Course No.: 132. Credits: 3. Title: General Psychology I. Contact: David B. Miller. Content Area: CA 3 Science and Technology
Department: PSYC Course No.: 132 Credits: 3 Title: General Psychology I. Contact: David B. Miller Content Area: CA 3 Science and Technology Catalog Copy: 132. General Psychology I Course Information: (a)
More informationOn-line scheduling algorithm for real-time multiprocessor systems with ACO
International Journal of Intelligent Information Systems 2015; 4(2-1): 13-17 Published online January 28, 2015 (http://www.sciencepublishinggroup.com/j/ijiis) doi: 10.11648/j.ijiis.s.2015040201.13 ISSN:
More informationThe Set of Candidate Models
1 The Set of Candidate Models Formulation of critical hypotheses and models to adequately reflect these hypotheses is conceptually more difficult than estimating the model parameters and their precision.
More informationCurrent requirements for a major (page 83 of current catalog)
To present the proposal of the Department of Mathematics and Computer Science for changing the requirements for a major in Computer Science, we first describe the current major requirements followed by
More informationand responding to the concerns of their constituents and of the general public in order to take advantage of new opportunities and to anticipate and
introduction For more than a century the business corporation has been a successful and widely adopted institutional arrangement for creating and distributing wealth. But the power and purpose of corporations
More informationIn developmental genomic regulatory interactions among genes, encoding transcription factors
JOURNAL OF COMPUTATIONAL BIOLOGY Volume 20, Number 6, 2013 # Mary Ann Liebert, Inc. Pp. 419 423 DOI: 10.1089/cmb.2012.0297 Research Articles A New Software Package for Predictive Gene Regulatory Network
More informationPsychology Learning Goals and Outcomes
Psychology Learning Goals and Outcomes UNDERGRADUATE PSYCHOLOGY MAJOR LEARNING GOALS AND OUTCOMES: A Report (March 2002) American Psychological Association, Task Force Members: Jane S. Halonen, Drew C.
More information5,6 2, 4, 6, 9, 10, 11, 15, 18, 19, 20 2, 16, 21 18, 19, 20, 22 14, 23
STATEMENTS FOR FIFRA SAP ON TERMITE BAIT PRODUCT PERFOMANCE TESTING GUIDELINES By Brian T. Forschler Department of Entomology College of Agricultural and Environmental Sciences University of Georgia Athens,
More information