Comparative Animal Behavior, Ethology, and Sociobiology COMMUNICATIONS

1. Comparative Animal Behavior, Ethology, and Sociobiology COMMUNICATIONS Neil GreenbergDepartment of Ecology and Evolutionary BiologyUniversity of Tennessee

2. What is communication?Classical definitions • “Any sharing of information” – Smith, 1984 • “… an action on the part of one organism (or cell) that alters the probability pattern of behavior in another organism (or cell) in a fashion adaptive to either one or both of the participants” – Wilson, 1975 • “The transmission of a signal from one animal to another such that the sender benefits, on average, from the response of the recipient” – Slater, 1983

3. Possible outcomes of communication Mutual benefit – true communication Sender benefits – manipulation/deceit Receiver benefits – eavesdropping Neither benefits – Highly unlikely

4. Levels of Communication Vegetative Tonic Phasic Signal Level (biosocial / psychosocial influences) Symbolic (develops through social experience) Language (abstract)

5. INPUTdetection of signals from the internal or external sense organs DEVELOPMENT (e.g., when in an animal’s life does signal detection (or lack thereof) occur; can it be influenced by experience?) (e.g., when sense organ is mature … but it might need environmental stimulation to do that!) ECOLOGY (e.g., can signal detection be influenced by the abiotic or biotic context) (e.g., when sense organ is mature … but it might need environmental stimulation to do that!) EVOLUTION (e.g., what capacities have been transmitted from prior generation(s) and what capacities will be passed on? (e.g., genes from ancestors, memes from ancestors, caregivers, peers) PHYSIOLOGY (e.g., what is the more-or-less proximate cause within the individual?)(e.g., when sense organ is mature … but it might need environmental stimulation to do that!)

6. What we know of nature is necessarily limited • CONSTRAINTS: our sense organs have evolved to detect only that which is essential to survival – to the meeting of our needs. The umwelt: there are many qualities of the world we have no “need” to know about – so the capacity to detect and act upon those qualities is weak at best and may never have evolved . . .

7. So our model of the world is necessarily fragmented and incomplete The ILLUSION of completeness is created by the mind so that we can take effective action rather than being indecisive. The illusion is perfected by (1) “extrapolating” by projecting information from within a “zone of confidence” to an outside area of less certainty or (2) “interpolating” or “filling in” from adjacent cells

8. we are a dynamic, endlessly shifting assemblage of individual attributes(think of hundreds of connected mobiles) In this environment, new fragments of information are “tested” for the best possible (not perfect) approximation of the real world (“correspondence”) and then assembled into the best possible (not perfect) story (“coherence”) • Life is the art of drawing sufficient conclusions from insufficient premises. • (Samuel Butler Notebooks (1912) ch. 1)

9. REALITY TESTING:does our inner model of the outer world correspond to reality? CORRESPONDENCE and COHERENCE

10. TEACHING is an act of COMMUNICATION • Teaching and learning involves “MEMES” – the cultural equivalent of biological units of information: “GENES.” • Art of Communication? • “...painting, that is to say the material thing called painting [is] no more than the pretext, than the bridge between the mind of the painter and that of the spectator.” (Eugene Delacroix 1850) • “It takes two to speak the truth-- one to speak, and another to hear.” (Thoreau1849)

11. EVOLUTION of BEHAVIOR “In animals, almost invariably, a change in behavior is the crucial factor initiating evolutionary innovation” (Ernst Mayr 1988). Behavior creates new selective pressures (Mark Baldwin via Deacon 1998)

12. The evolution of communicative signalsRITUALIZATION There is often “selection pressure” for social signals to become more precise, less ambiguous . . . Termed “ritualization” it is of great intrinsic interest to the student of social behavior … BUT the processes observed also exemplify the ways in which units of behavior can be transformed in function over time – evolve. Any behavioral pattern is a candidate for ritualization if it can communicate information Many behavioral patterns “multitask”, particularly those that communicate information about the internal state that might affects behavior

13. The evolution of communicative signalsRITUALIZATION MOTOR PATTERNS (“somatic”) • Intention movement of (e.g.) body, limbs, ears, tail • Ambivalent Posture or movement . . . raising or lowering head (e.g., dominance) AUTONOMIC REFLEXES • Alimentary: Increase or decrease in salivation. Sphincter control, urination, defecation. (e.g., territorial marking) • Circulatory: Pallor, flushing, vasodilation of sex organs. Fainting.(e.g., skin patches) • Respiratory: Changes in respiratory rate or amplitude. Gasping, sighing, panting. (e.g., inflation displays, hissing, speech (?) • Thermoregulatory: Sweating, pilomotor responses. (e.g., hair or feather erection, scent signals) Electrodermal response • Lacrimatory: weeping.

14. The evolution of communicative signalsRITUALIZATION Ways in which the displays can become elaborated(Morris 1966): 1. Development of conspicuous structures 2. Schematization("simplification") by means of (e.g.) • Lowering of threshold changes • Rhythmic repetition • Exaggeration or omission of specific components of a movement • Change in speed or vigor of a movement When the movement comes to be governed by causal factors other than those that governed the source of the display it is said to be emancipated.

15. The Peacock’s Tail • raised by feather pilomotor muscles • an ancient autonomic thermoregulatory mechanism • Ordinarily hidden • displayed when aroused

16. The Lizard’s Flag • Effected by the hyoid apparatus • An ancient mechanism activated by stress • Ordinarily hidden • displayed when aroused